Professional Documents
Culture Documents
Durio
Durio
Durio
MICHAEL J. BROWN
Department of Plant Science
MacDonald College, McGill University,
Quebec, Canada
This publication has been funded by the International Plant Genetic Resources Institute.
Citation
Michael J. Brown. 1997. Durio - A Bibliographic Review (R.K. Arora, V. Ramanatha
Rao and A.N. Rao, Editors). IPGRI office for South Asia, New Delhi.
iv
Preface
My desire to write this book grew out of an innate and deeply rooted
concern to understand the durian. In the late 1980s, a vigorous search of
scientific abstracts led me to believe that a bibliography on the subject would
only fill 1 or 2 type-written pages; the idea that a bibliographic review
resulting in a book would be necessary to fully examine the subject was
unthinkable. Today, I would estimate that approximately 1200 research
articles, book chapters, conference abstracts etc. have been produced which
pertain to durian. At the current rate of increase, this figure will very likely
double over the next 20 years. Thus, we stand at a crossroad. If our future
researches are really to advance our knowledge forward, we need to come to
terms with what we already know.
V
In this work, great pains have been taken to give full credit to all the
authors. Much time and effort have been devoted to obtaining original research
articles to confirm statements and ideas presented in more general reviews. In
cases where original results were published using the Imperial system of
measurements, a metric equivalent is presented parenthetically. Where research
on species, that are no longer taxonomically recognized, is discussed, parenthetic
notes are given to help clarify their identity.
I can in no way take full credit for the massive amount of effort that
went into compiling the bibliography. I wish to extend thanks to the numerous
libraries and institutions without whose aid in supplying original works and/
or photocopies, this book would not have been possible: University of Guelph
Library, Canada; Universiti Pertanian Malaysia Library; National Library of
Malaysia; Malaysian National Agricultural Library; Management Information
Systems Division, MARDI, Malaysia; University of Los Banbs Library,
Philippines; Kasetsart University Library, Thailand; Thailand Institute of Scientific
and Technological Research; Hunt Botanical Library, Pennsylvania; The Library
of the Herbarium Universitatis Florentinae, Italy; The Library of the New
York Botanical Gardens; The Library of the Jardin Botanique National de
Belgique, Belgium; The British Library (Oriental and India Office Collections);
Centre for Scientific Documentation and Information, Indonesia; SEAMEO-
BIOTROP, Bogor, Indonesia; Forest and Nature Conservation Research and
Development Centre, Bogor, Indonesia; Library of The Royal Botanic Gardens,
Kew, England; Lloyd Library and Museum, Cincinnati USA; and The Solomon
Islands National Library.
I am grateful to the many authors who supplied reprints of their
research articles. I am indebted to the friendship and excellent library skills
of Mr. David Bantroch, which greatly enhanced the scope and content of the
bibliography. I am also indebted to those who graciously supplied or helped
obtain copies of articles from different locations across the world: Dr. Gordon
Brown, Carol Bowes, Paul d’Amboise, Peter Toorop and Dr. Andrew Powell.
I also express my gratitude to the International Plant Genetic Resources Institute
for supporting the publication of this work, and to Drs. R.K. Arora, V. Ramanatha
Rao and A.N. Rao for editing the manuscript.
Finally, I would like to acknowledge the invaluable assistance of
those who helped translate articles or parts thereof: Nusin Brown for translation
of Turkish items; Aldo De Moor, Dr. Annette Nassuth and Peter Toorop for
translation of Dutch and German items; Dr. Wataru Mitsuhashi for the translation
of Japanese items, Chumnum Wongmanee for translation of Thai articles and
my wife Nathalie Bourgouin-Brown for translation of several French documents.
Michael J. Brown
vi
Contents
Foreword iii
Preface V
Contents vii
Introduction
Taxonomic History 2
Origin of the word durian; Durian poetry; Historical works;
Authority for Durio zibethinus; Formation of the modern
concept of Bombacaceae; Wild species of durian; Post
Kostermans (1958b) treatment; Keys to the species; Future
taxonomic work
Morphology 22
Durian theory; Floral morphology; Pollen morphology; Fruit
morphology; Fruit teratology; Other teratologies; Ovule and
seed morphology and development; Leaves; Roots; Tree
architecture; Chromosome number
Edibility, Composition and Uses of the Fruit 34
Edibility; Nutritive constituents; Fatty acids; Smell;
Miscellaneous uses of fruit
Medicinal and Toxicological Properties 51
Durians and alcohol; Medicinal properties; Febrifugal and
anti-malarial properties; Vermifugal properties; Treatment
of jaundice; Diabetes; Aphrodisiac; Miscellaneous medicinal
properties
Seeds 56
Mature seed constituents; Culture of seeds and seed size;
Viability; Germination
Pollination Biology 61
Anthesis; Natural pollinators; Early ovary abscission vs
premature fruit drop; Self-incompatibility; Heterostyly;
Mechanisms of self-incompatibility; Empirical evidence;
Premature fruit drop; Manipulating premature fruit drop;
Leaf flushing; Species which feed upon durians; Natural
dispersal of seeds
Fruiting Seasons 69
Maturation of buds; Environmental effects on flowering;
vii
Manipulating seasonality
Ecology, Origin . and Distribution 72
Centre of diversity; Wild form of Durio zibethinus; Attempts
at introduction
Clones 77
Clonal selection and hybridization; Clonal identification
Nursery Care and Cultivation 81
Seeds; Branch pruning; Root pruning; Application of fertilizer;
Soil conditions; Water relations; Transplanting; Intercropping
Post-harvest Technology
Grading; Shipping and cold storage of fruit; Packaging; 87
Ripening of fruits; Effects of atmosphere; Plant growth
regulators; Post-harvest technology; Processed food products
and their packaging
Forestry Aspects 94
Timber characteristics of durio species and close relatives;
Boschia griffithii Mast. [=D. griffithii (Mast.) Bakh.];
Coelostegia griffithii Mast.; Cullenia excelsa Wight; Durio
carinatus Mast.; Durio dulcis Becc.; Durio kutejensis (Hassk.)
Becc.; Durio ?lowianus Scort. ex King; Durio malaccensis
Planch. ex Mast.; Durio ?oblongus Mast.; Durio oxleyanus
Griff.; Durio testudinarum Becc.; Durio zibethinus L.,
Durability of ‘Durian’ timber; Uses of ‘Durian’ timber;
Properties of ‘Durian’ timber; Wood anatomy
Major Diseases, Parasitism and Associated Organisms 100
Bacteria; Fungi; Lichens; Algae; Ferns; Angiosperms; Insects;
Nematodes; Other animals; Hyperparasitism in durians
Vegetative Propagation of Durians 111
Etiolated shoot method; Double root system; Approach grafting;
Inarching; Top grafting; Budding techniques; Other asexual
propagation techniques; Grafting to other species; Advanced
planting material; ‘Hybrid’ Durians
Economics and Prospects for Development 117
Genetic Resources and Conservation 120
Bibliography 126
viii
Durio — A Bibliographic Review
Introduction
The genus Durio is native to South East Asia with its centre of
diversity in Borneo (Mendoza 1941; Lim 1990). The genus comprises
approximately 30 known species, of which only Durio zibethinus is cultivated
for its fruits to a great extent (Lim 1990). As a rain-forest tree, it typically
attains heights of 30-40 m (Tidbury 1976) and diameters of 2-2.5 m, but the
cultivated varieties in an orchard, especially when grafted, grow no higher
than 12 m (Malo and Martin 1979). Although relatively unknown to the
western world, the durian is a valuable commodity in South East Asia, and
has had a profound effect on the history and culture of that part of the world.
James Low (1836) recorded that the king of Ava had fruits transported to him
at Amerapoora, ‘by relays of horsemen, and by boats pulled by 40 or 50
men’. The durian fruit's reputation precedes it wherever it goes: ‘durian is to
fruit what limburger is to cheese and pornography is to literature’ (Anon.
1979a).
Although little research has been carried out in the past on durian,
this trend is slowly starting to change as plans to improve the quality and
consistency of durian fruits develop. This is evidenced by the success of the
recent release of the first ‘hybrid’ durians in Malaysia (Othman 1991).
Taxonomic History
The durian has influenced various cultures of South East Asia for
millennia, but has only been known to the western world for about 600 years.
In this section, attempt is made to trace the scientific description of durian
from its earliest origins to the present day. Early descriptions and knowledge
were largely of a morphological and taxonomic nature. Although accounts on
the effects of the fruit on human physiology abounded, our knowledge of
these have expanded enormously in the last few decades and thus this aspect
will be discussed in a separate section. Tracing the origins of durian research
to their beginnings not only allows us to place certain information in proper
historical perspective, but also an understanding of how the current taxonomic
state of affairs, and perhaps much of the confusion, has arisen.
Origin of the word durian : The word durian (Durio) without doubt originates
from the Malay word ‘duri’ which means spine (Don 1831). The word ‘zibethinus’
is a reference to the Indian civet cat Viverra zibetha. Don (1831) provided the
following information: ‘the fruit is used as bait to entrap the civet-cat, which
is very fond of it; hence the specific name’. Others have suggested that zibethinus
refers to the smell of the fruit which is it’s (and the civet cat’s) most legendary
characteristic (Hawson 1983; Watson 1984). The accuracy of this comparison
is perhaps best summed up by Barrett (1912). ‘According to the specific name
zibethinus, the fruit should osphresiologically remind one of the civet cat; the
writer, however, after having seen and smelled live civets in Mozambique,
does not concur in this idea.’ Nevertheless, the fruit is occasionally referred
to as the civet cat fruit (Gamble 1881; Watt 1890; MacMillan 1909, 1912;
Anon. 1952; Singh et al. 1983). The Latin name was coined by Linnaeus who
apparently never encountered an actual specimen of a durian, and based his
description entirely upon that provided in the ‘Herbarium Amboinense’ (König
1804). As Rumphius (1741) refers to the use of durian fruits to catch civet
cats in his ‘Herbarium Amboinense’, it seems likely that this was the reason
for the name. De Candolle (1824) also states that the name arises from the
fact that civets eat durians.
suggests that the conservation of the word ‘durian’ in different native Indonesian
languages probably indicates its early spread by the Malays.
Many authors give Malay vernacular names for the various species.
Malay names can often be quite useful in aiding identification (Corner 1988);
the taxonomy of many of the wild species of Durio is still in some disarray,
and it is quite likely that the Malay names are more reliable in some instances.
By far the most comprehensive, and probably most accurate, list of vernacular
names for different species of durian are those given by Kostermans (1958b).
Durian poetry : For whatever reason, the durian has appealed to the artistic
side of people for a long time. Many Malay idioms contain references to
durians (Kostermans 1958b). There are also several published poems pertaining
to durians (Whiteside 1914; Slate 1974; Chin 1979; Chin 1980a; Bantroch,
1995). A short story simply entitled ‘Durian’ by A.R. Roces (1949) delightfully
conveys the not-so-subtle nature of durian fruits to the reader.
edn. 13, 581 (1774)]’ is the earliest described species. They are both correct.
They differ in that Ridley (1922) attributes this work to Linnaeus, while
Wyatt-Smith (1953a) attributes the same work to Murray.
1
According to what is unquestionably the most thorough source of information on taxonomic
literature, "the botanical novelties in this [13th] edition still stem from Linnaeus and must be
attributed to him. Murray acted here as editor" (F. Stafleau and R.S. Cowan 1981), Taxonomic
Literature. 2nd edition. Volume 3). Yet, Farr et al. (1979) in the ‘Index Nominum Genericorum’,
who attempted, after indepth research, to compile a list of valid plant genera, type species and
their authorities, cites D. zibethinus Murray (Syst. Veg. edn. 13:581. 1774) as the type.
Murray was unquestionably the editor of the 13th edition of ‘Systema Vegetabilium’;
Murray received a copy of the manuscript for the 13th edition from Linnaeus in 1771, who
asked him to find a publisher for it in Germany (H. Goerke 1976), Linnaeus and the Murray
family, Taxon 25(1), 15-19). Although Murray was awarded the status of Editor, his function
was only to obtain a publisher for this work. Accordingly, the title page bears the Latin
inscription "Accessionibus et emendationibus novissinus manu perillustris auctoris scriptis" .
Thus, the common durian is correctly referred to Durio zibethinus L.
Durio — A Bibliographic Review 7
Numerous species and varieties of durian have been named over the
years, over half of which are no longer recognized. Despite this, several of
these ‘defunct’ names continue to be used. This is hardly surprising considering
the complexity surrounding the taxonomy. Table 1 attempts to present all the
published names of species of Durio and their synonyms. The authority(ies)
of all published names along with the date of publication have been listed.
Where name changes have occurred, details are provided as far as possible.
The actual details of the current composition of several species (with respect
to particular herbarium specimens) is far more complex than can actually be
Durio — A Bibliographic Review 10
tabulated. The relevant works for further details should be consulted. The
genus Cullenia is also included as it has, in part, been previously included
within the genus Durio and may well be again. As diagrams have, and still
do have, a large role to play in the naming and identification of species, a list
of sources for figures of plant parts for currently recognized species is presented
in Table 2.
(Contd....)
Durio — A Bibliographic Review 11
Table 1. Contd.
-is a mere synonym of D. ceylanicus Gardn. (sensu Kostermans 1958b)
-split among C. ceylanica, C. rosayroana and C. exarillata (sensu Robyns
1970)
C. rosayroana Kosterm. (1956)
-includes part of C. excelsa Wight. (sensu Kostermans 1956)
C. zeylanica (Gardn.) Wight ex K. Schum. (1895)
-transferred to D. ceylanicus Gardn. (sensu Bakhuizen Van Den Brink
1924a)
Durio acuminatissima [Kostermans and Soegeng-Reksodihardjo 1958]
=D. acuminatissimus Merr.
D. acuminatissimus Merr. (1924)
-included in D. zibethinus L. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. acutiminatissimus Merr. [Lim 1990]
=D. acuminatissimus Merr.
D. acutifolia (Mast.) Kosterm. (1953b)
-now included in D. acutifolius (Mast.) Kosterm. (sensu Kostermans
and Soegeng-Reksodihardjo 1958)
D. acutifolius (Mast.) Wyatt-Smith (1953a)
-includes B. acutifolia (Mast.) (sensu Wyatt-Smith 1953a)
-includes D. griffithii (Mast.) Bakh. var. acutifolius (Mast.) Bakh. (sensu
Wyatt-Smith 1953a)
D. acutifolius (Mast.) Kosterm. & Soegeng. (1958)
-includes D. acutifolius (Mast.) Wyatt-Smith
-includes B. oblongifolia Ridl.
-includes D. griffithii (Mast.) Bakh. var. acutifolius (Mast.) Bakh.
D. affinis Becc. (1889)
-includes D. malaccensis Planch. ex Mast. (sensu Kostermans and
Soegeng-Reksodihardjo 1958)
D. beccarianus Kosterm. & Soegeng. (1958)
D. bukitrayaensis Kosterm. (1990)
D. burmanicus Soegeng. (1965)
D. carinatus Mast. (1874b)
-includes D. cupreus Ridl. (sensu Kostermans and Soegeng-Reksodihardjo
1958)
(Contd....)
Durio — A Bibliographic Review 12
Table 1. Contd.
D. carinatus Mast. var. bintulensis Becc. (1889)
D. ceylanica Gardn. [Wight 1852]
= D. ceylanicus Gardn.
D. ceylanicus Gardn. (1847)
-includes D. zibethinus Moon (sensu Gardner 1847)
-is now C. ceylanica (Gardn.) K. Schum. (sensu Robyns 1970)
D. conicus Becc. (1889)
-is now D. dulcis Becc. (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. crassipes Kosterm. & Soegeng. (1958)
D. cupreus Ridl. (1938)
-included in D. carinatus Mast. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
-re-erected to species status (Kostermans 1961)
D. dulcis Becc. (1886)
-reduced to D. conicus Becc. (sensu Bakhuizen Van Den Brink 1924a)
-synonym of D. graveolens (sensu Wyatt-Smith 1953a)
-includes D. oblongus Mast. (sensu Kostermans & Soegeng-Reksodihardjo
1958)
-includes D. conicus Becc. (sensu Kostermans & Soegeng-Reksodihardjo
1958)
D. excelsus (Korth.) Bakh. (1924a)
-includes part of D. excelsus (Korth.) Bakh. var. typicus Bakh. (sensu
Kostermans and Soegeng-Reksodihardjo 1958)
-includes D. griffithii (Mast.) Bakh. var. heteropyxis (Griff.) Bakh.
(sensu Kostermans and Soegeng-Reksodihardjo 1958)
-includes most of B. excelsa Korth. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. excelsus (Korth.) Bakh. var. typicus Bakh. (1924b)
-part is now D. excelsus (Korth.) Bakh.; other part is D. griffithii
(Mast.) Bakh. (sensu Kostermans and Soegeng-Reksodihardjo 1958)
D. excelsus (Korth.) Bakh. var. grandiflorus (Becc.) Bakh. (1924b)
-transferred to D. grandiflorus (Mast.) Kosterm. & Soegeng. (sensu
Kostermans and Soegeng-Reksodihardjo 1958)
(Contd....)
Durio — A Bibliographic Review 13
Table 1. Contd.
D. falcata [Stadelman 1966]
-name of no taxonomic standing
D. foetida Thunb. (1796)
-name of no taxonomic standing, synonym of D. zibethinus L.
D. grandiflorus (Mast.) Kosterm. & Soegeng. (1958)
-includes part of B. grandiflora Mast. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
-includes B. excelsa Korth. [sensu Merrill 1921, 1929] (sensu Kostermans
and Soegeng-Reksodihardjo 1958)
-includes D. excelsus (Korth.) Bakh. var. grandiflorus (Becc.) Bakh.
(sensu Kostermans and Soegeng-Reksodihardjo 1958)
D. gratissimus Becc. (1889)
-is now included in D. oxleyanus Griff. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. graveolens Becc. (1889)
-reduced to D. conicus Becc. (sensu Bakhuizen Van Den Brink 1924a)
-includes D. dulcis (sensu Wyatt-Smith 1953a) [this inclusion not valid
under Kostermans and Soegeng-Reksodihardjo 1958]
-re-erected to D. graveolens Becc. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. griffithii (Mast.) Bakh. (1924a)
-includes some of D. griffithii (Mast.) Bakh. var. heteropyxis (Griff.)
Bakh. (sensu Kostermans and Soegeng-Reksodihardjo 1958)
-includes some of D. excelsus (Korth.) Bakh. var. typicus Bakh. (sensu
Kostermans and Soegeng-Reksodihardjo 1958)
D. griffithii (Mast.) Bakh. var. heteropyxis (Griff.) Bakh. (1924b)
-a synonym of D. excelsus (Korth.) Bakh. with the exception of a
Sumatra specimen (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. griffithii (Mast.) Bakh. var. acutifolius (Mast.) Bakh. (1924b)
-included under D. acutifolius (Mast.) Kosterm. & Soegeng. (sensu
Kostermans and Soegeng-Reksodihardjo 1958)
D. kinabaluensis Kosterm. & Soegeng. (1958)
-includes D. kutejensis (Hassk.) Becc. forma kinabaluensis Bakh.
(Contd....)
Durio — A Bibliographic Review 14
Table 1. Contd.
D. kutejensis (Hassk.) Becc. (1889)
-includes L. kutejensis Hassk. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. kutejensis (Hassk.) Becc. forma kinabaluensis Bakh. [Wyatt-Smith 1953a]
-is now D. kinabaluensis (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. lanceolatus Mast. (1874b)
-reduced to D. singaporensis Ridl. (sensu Wyatt-Smith 1953a)
-re-erected to species status (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. lissocarpus Mast. (1874b)
-reduced to D. carinatus Mast. (sensu Bakhuizen Van Den Brink 1924a;
Wyatt-Smith 1953a)
-re-erected to species status (sensu Kostermans and Soegeng-Reksodihardjo
1958)
D. lowianus Scort. ex King (1891)
-includes D. wrayii King. (sensu Kostermans 1958a)
-includes D. zibethinus L. var. roseiflorus Corner (sensu Kostermans
1958a)
D. lowii Hook. [Sutisna and Soeyatman 1985]
-name of no taxonomic standing, possibly D. lowianus Scort. ex
King
D. macrantha Kosterm. (1992a)
D. macrolepis Kosterm. (1958a)
-specimens fractured into D. pinangianus (Becc.) Ridl. and D.
macrophyllus (King) Ridl. (sensu Kochummen et al. 1970)
D. macrophyllus (King) Ridl. (1922)
-reduced to D. oblongus Mast. (sensu Bakhuizen Van Den Brink 1924a)
-re-erected and includes D testudinarum Becc. var. macrophylla King
(sensu Kostermans 1958a)
D. malaccensis Planch. ex Mast. (1874a)
-includes D. sumatranus Becc. (sensu Kostermans 1958a)
-part has been removed to D. affinis Becc. (sensu Kostermans and
Soegeng-Reksodihardjo 1958)
D. mansoni (Gamble) Bakh. (1924b)
-includes B. mansoni Gamble (sensu Kostermans 1958a)
(Contd....)
Durio — A Bibliographic Review 15
Table 1. Contd.
D. oblongus Mast. (1874b)
D. oxleyanus Griff. (1845)
-includes D. gratissimus Becc. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
-includes D. griffithii Planch. ex King. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
D. perakensis King (1891)
-a name of no taxonomic standing, possibly D. lowianus Scort. ex
King
D. penangianus [Kochummen and Wyatt-Smith 1979]
=D. pinangianus (Becc.) Ridl.
D. pinangianus (Becc.) Ridl. (1922)
-includes D. testudinarum Becc. var. pinangianus Becc. (sensu Kostermans
1958a)
-includes some of D. macrolepis Kosterm. (sensu Kochummen et al.
1970)
D. purpureus Kosterm. & Soegeng. (1958)
D. singaporansis Ridl. [Lim 1990]
=D. singaporenis Ridl.
D. singaporensis Ridl. (1922)
-reduced to D. oblongus Mast. (sensu Bakhuizen Van Den Brink 1924a)
-re-erected by Wyatt-Smith (1953a) as a possible synonym of D.
sumatranus
-retained as D. singaporensis Ridl. (sensu Kostermans 1958a)
D. singapurensis [Corner 1978]
=D. singaporensis Ridl.
D. spontaneus Bakh. [Van Steenis 1949]
-is now D. lowianus Scort. ex King. (sensu Kostermans 1958a)
D. stercoraceus Noronha (1790)
-synonym of D. zibethinus L.
D. sumatranus Becc. (1889)
-may include D. singaporensis Ridl. (sensu Wyatt-Smith 1953a)
-synonym of D. malaccensis Planch. ex Mast. (sensu Kostermans
1958a)
(Contd....)
Durio — A Bibliographic Review 16
Table 1. Contd.
D. testudinarium Becc. [Lim 1990]
=D. testudinarum Becc.
D. testudinarum Becc. (1889)
D. testudinarum Becc. var. pinangianus Becc. (1889)
-now D. pinangianus (Becc.) Ridl. (sensu Wyatt-Smith 1953a and
Kostermans 1958a)
D. testudinarum Becc. var. macrophylla King (1891)
-lifted to D. macrophyllus (King) Ridl. (sensu Wyatt-Smith 1953a and
Kostermans 1958a)
D. testudinarum Becc. var. macrophyllus King [Corner 1939]
=D. testudinarum Becc. var. macrophylla King
D. wrayi [Ridley, 1922; Kostermans and Soegeng-Reksodihardjo 1958]
=D. wrayii King
D. wrayii King (1891)
-reduced to D. lowianus Scort. ex King (sensu Kostermans 1958a)
D. wyatt-smithii Kosterm. (1958a)
D. zeylanica [Worthington 1959]
=D. ceylanicus Gardn.
D. zibethianus Murr. [Kanehira 1935]
=D. zibethinus L.
D. zibethinus L. (1774)
D. zibethinus L. var. roseiflorus Corner (1939)
-now D. lowianus Scort. ex King (sensu Kostermans 1958a)
D. zibethinus Moon (1824)
-synonym of D. ceylanicus Gardn.
D. zibethinus Murr. (1774) improper attribution of authority = D. zibethinus
L. (1774)
D. sp. “A” [Kochummen 1972]
-a new, as yet un-named and incompletely known species with similarities
to D. lanceolatus and D. kutejensis
Lahia kutejensis Hassk. (1844)
-is now Durio kutejensis (Hassk.) Becc. (sensu Kostermans and Soegeng-
Reksodihardjo 1958)
(Contd....)
Durio — A Bibliographic Review 17
Table 1. Contd.
Neesia griffithii Planch. ex King
-is now D. oxleyanus Griff. (sensu Kostermans and Soegeng-Reksodihardjo
1958)
Currently valid species are in bold; species of dubious, controversial or unknown status are in
italics, invalid species are in normal type. Dates in round brackets ( ) after a species name
represent the date of publication of the species by the preceding authority. Authors and dates
in square brackets [ ] represent publications in which an invalid or unrecognized name was cited
either by typographical error or erroneously, or cases where the author of the first published
description is not the authority.
Table 2. Sources for figures of currently valid species of Durio and close
allies
Table 2. Contd.
Table 2. Contd.
Kostermansia BW32
malayana
[Published diagrams of durian species: BW=black and white photo, C=colour
photo, LN=line drawing. Citations with 2 superscripts indicate the same figure
appears in two separate publications. An * indicates that additional clarifying
information is given in the following list of sources.]
5
Beddome (1869).
6
Cockburn (1976).
7
De Vogel (1980) *Note: Diagrams are of seedlings with attached seeds.
8
Foxworthy (1927).
9
Griffith (1854a) Plate 596
10
Kadambi (1954).
11
Köing (1804).
12
Korthals (1839-1842) *Note: What is depicted as Boschia excelsa is now interpreted by
Kostermans (1958b) as D. excelsus (Korth.) Bakh.
13
Kostermans (1956), 14Kostermans (1953b), 15Kostermans (1958a), 16Kostermans (1958b).
17
Kostermans (1958c), 18Kostermans (1990), 19Kostermans (1992a), 20Kostermans (1992b).
21
Lamarck (1823).
22
Meijer (1969).
23
Ochse (1927).
24
Ridley (1922) *Note: Part of the figure representing D. malaccensis was copied from Masters
(1874b). The copied parts include the anther at the top right, and the two bottom left
diagrams (longitudinal section of an ovary, and a spine bearing a peltate scale). According
to Kostermans (1958b), of these 3 copied diagrams, only the anther belongs to D.
malaccensis Planch. ex Mast.
25
Robyns (1970).
26
Setiadi (1991) *Note: The photo of D. testudinarum is captioned ‘Durian kura-kura’.
27
Soegeng-Reksodihardjo (1962), 28Soegeng-Reksodihardjo (1965).
29
Sprague (1915).
30
Stanton (1966).
31
Wettstein (1935) *Note: Figure 3 on page 806, showing the flower of D. affinis, is copied
from volume III, Table 24 of Beccari’s Malesia of 1889. This diagram was also copied
by Kostermans 1958b, appearing as part of Fig. 28.
32
Whitmore (1990).
33
Wight (1852) *Note: This figure is titled Cullenia excelsa Wight. which, according to Kostermans
(1958b), is equivalent to Durio excelsa Gardn.=Cullenia ceylanica (Gardn.) K.Schum.
However, Kostermans (1956) and Raizada (1957) consider Wight’s figures to be drawn
from a mixture of specimens of C. ceylanica and C. rosayroana, most of them being
C. rosayroana with the exception of the fruit depicted in figures 14-16 which are
probably C. ceylanica. More recently, Robyns (1970) has described a new species
Cullenia exarillata A. Robyns. which he claims is what is depicted in Wight’s original
figures, the interpretation of Kostermans (1956) and Raizada (1957) being in this regard
erroneous.
34
Winkler (1905).
35
Worthington (1959).
Durio — A Bibliographic Review 21
Keys to the species : As numerous new species have been described in the
last 50 years, and major reworking of species have also occurred, keys prior
to Kostermans (1958b) are now of little use. An exception to this might be
Wyatt-Smith (1953a), whose key to 19 species, based mainly on floral
characteristics, is largely in agreement with the classification of Kostermans
(1958b). Kostermans (1958b) presents a key in his monograph based largely
on floral characteristics. Soegeng-Reksodihardjo (1962) provided keys to six
edible species based on floral, fruit or vegetative characteristics.
Future taxonomic work : Despite all the taxonomic work which exists on
the genus Durio, it is quite clear that much remains to be clarified. Some
of the wild species are known only from very little and incomplete herbarium
specimens. For example, D. crassipes Kosterm. & Soegeng. is only known
from one herbarium specimen, consisting of a few flowers which are missing
the epicalyx (Kostermans 1958b). Some species have not been collected in
many years and may have already been extinct. An examination of the various
treatments of the genus, which have been published, reveals that the major
cause of the collapse of old species and the erection of new ones is the
subjective decision as to how physically different two herbarium specimens
have to be in order to be given different names. Further, monographing of
species in the style so far established for them seems unlikely to provide
more insight on the matter. It is difficult to imagine how such further
taxonomic shuffling can really address the most pressing and meaningful
questions that are in need of answers. Future work involving proper cladistical
analysis of as many characters as can be obtained from the relevant herbarium
(and other) specimens, perhaps coupled with RFLP (restriction fragment length
polymorphism) mapping and isozyme analysis would be valuable. Isozymes
have already proved useful in distinguishing clones of D. zibethinus (Salma
1993). It has also been suggested that the position and morphology of the
leaf trichomes may also be of taxonomic value (Salma 1991). Furthermore,
chromosome counts might be of use in addressing taxonomic questions
surrounding durian. Many of the species of Durio are known from living
specimens, thus crossing experiments are possible. The structure of many
durian flowers lends itself to artificial pollination, and the life span of durian
pollen has been demonstrated to be sufficiently long under appropriate
conditions to allow such crossing. Some crossing experiments have been
conducted and offer the exciting prospect of improved trees, especially with
regard to disease resistance.
Durio — A Bibliographic Review 22
Morphology
A fair amount of morphological and anatomical information on durians
has been published. The majority of this information deals with the structure
of the flowers and fruits, but some information is available on the leaves,
roots, wood anatomy and seeds. One of the most fascinating developments
from the study of the morphology of durian fruits has been E.J.H. Corner’s
durian theory (Corner 1949). A full examination of this theory is well beyond
the scope of this work, as much of it involves species extraneous to the topic,
but as this theory is forever linked to the durian fruit, it will be briefly
discussed. Additionally, the tree architecture of durian has been analyzed. The
anatomy of the wood will be discussed in a later chapter in relation to the use
of durian in forestry, as this is the field in which such information is most
useful. As chromosome counts have often been used in conjunction with
taxonomic and morphological analysis, this information is also included under
this section.
Durian theory : In 1949, Corner elaborated his ‘Durian Theory’ which, among
other things, predicted that the ancestral angiospermous fruit was large, spiny
and dehiscent, bearing large seeds covered in colourful fleshy arils. This type
of fruit is more or less typified by that of the durian tree (Durio zibethinus L.).
This theory was spurred by his observations of species with strikingly similar
fruit morphologies in numerous unrelated angiosperm families. These species
are usually very rare, and their fruits atypical of the other more common
members of the family. Through a series of arguments, he proposed it was
unlikely that this peculiar fruit type had evolved independently numerous
times, and more likely that it represents a relic. Several arguments have been
raised against this theory, most notably by Parkin (1953) and Van Der Pijl
(1952, 1955). The main points of contention are the subjectivity of Corner’s
observations, and the denial that rarity necessarily represents antiquity. An
attempt to refute this theory was published by Datta and Biswas (1969), but
their ‘argument’ was based on an obvious misunderstanding of Corner’s theory,
and thus provides neither support nor evidence against it. Corner has since
expanded upon his theory (Corner 1953, 1954a, 1954b), and it has been given
some support by others (Mabberley, 1974a,b; Von Teichman and Van Wyk
1991, 1994). The most objective and perhaps most useful new evidence comes
from Von Teichman and Van Wyk (1991) whose use of statistical character
associations revealed that ‘durian fruit and seed-like’ characteristics (i.e.,
recalcitrance, arils, pachychalazy, etc.) were significantly correlated with the
occurrence of other character traits generally regarded as primitive. More
recently, Von Teichman and Van Wyk (1994) have elaborated on the idea
hinted at by Corner (1949) that recalcitrance (high seed moisture content and
Durio — A Bibliographic Review 23
a short period of seed viability) is the ancestral state of seeds, orthodoxy (the
ability to withstand desiccation) having evolved later under selection pressure.
This avenue of investigation suggests that recalcitrant seeds may share
morphological and physiological commonalities by descent. This concept may
be of substantial importance in the understanding of recalcitrant seeds and
definitely opens new lines of enquiry. However, to this date, the validity of
Corner’s durian theory remains in dispute, and its domain of applicability is
yet to be firmly established.
or less into a tube. The anthers are unilocular, midfixed, strongly recurved
and twisted, and dehisce via a single slit (Croft 1981). According to Davis and
Bhattacharya (1974), the anthers are two-celled or occasionally one-celled.
An extremely detailed examination of the stamens of several species of Durio
is presented by Van Heel (1966), and the development of the anther-wall has
been described in detail by Soepadmo and Eow (1977).
Figure 2. An open flower from D24 durian and its parts. The flower was
collected in the evening of anthesis, the male and sterile floral
organs abscind during the night. Figures B-E represent parts collected
the next morning.
A) A mature flower, projecting from the epicalyx which normally
splits into 2 or 3 sections. The sepals are fused to form a short tube
with a swollen base. The 5 petals emerge through the calyx tube
and reflex, exposing the numerous filaments and the stigma.
B) The gynoecium of the flower. All of the floral parts except the
gynoecium have abscinded. The ovary is covered in peltate scales,
the long style is often kinked in flowers from this particular clone.
The stigma is also visible. The receptacle bears visible scars left
from the abscision of the other floral parts.
C) The calyx of the flower. The fused sepals form a swollen base
which holds the nectar produced by nectaries inside the calyx.
D) A staminal phalange consisting of 7 filaments. The filaments
are connate at the base. Note that the filaments near the centre of
the phalange are longer than those at the sides.
E) A petal collected from under the tree the morning after anthesis.
Note that the petals are reflexed to expose the numerous filaments
enclosed within them.
Durio — A Bibliographic Review 26
Pollen morphology : The pollen grains of durian were probably first described
by Van Der Pijl (1936) who described them as 90 mm in diameter (wet), and
covered in a sticky mass which stains red with Sudan III. The pollen grains
are approximately spherical (Soepadmo and Eow 1977). Davis and Bhattacharya
(1974) measured pollen grain size from left and right twisted flowers, and
discovered that left twisted flowers had slightly, though consistently, larger
pollen grains from those of right twisted flowers (see also footnote 1). According
to Soepadmo and Eow (1977), the pollen grains of Durio zibethinus are
approximately 80-150 µm in diameter, 3-4 or rarely 6 porate with a smooth
and sticky exine. Davis and Bhattacharya (1974) calculated durian pollen
grains to vary from 20 to 80 µm with a mean of approximately 55 µm for dry
pollen and 67 µm for water soaked pollen. Erdtman (1972) states that the
equatorial diameter is 50-100 µm. A study by Salakpetch et al. (1992) included
the measurements of the equatorial and polar axes of pollen grains from four
Thai cultivars. This study showed that there were slight differences between
cultivars, however, the average length of both axes of all cultivars was of the
order of 15 µm. I have no explanation for the much smaller dimensions
provided by this study compared with those mentioned previously.
3
‘Kepala’ is Malay for head, while ‘gajah’ is the Malay term for the Indian Elephant ( Elephas
maximus).
Durio — A Bibliographic Review 28
useful for characterizing durians into distinct groups, and both are highly
heritable characters.
Rao and Singh (1964) reported that occasionally the floral meristem
continued to be active after formation of all the regular floral parts resulting
in the production of stamens and carpels within the ovary on the central axis
of the fruit. The anthers were found to contain pollen, but none of the carpels
contained mature ovules. No petals or sepals were seen to be produced. The
authors noted that these anomalous structures do not take the place of previous
structures, but are superfluous in nature.
‘A fruit of which the margins of the carpels with their warts and
hunches at the inner-side interlock like a cog-wheel’ was described by Costerus
and Smith (1932).
Other teratologies : Apart from fruits occurring within fruits, petaloid stamens
have also been described in durian (Winkler 1905). Soepadmo and Eow (1977)
related that they had twice observed the formation of binucellate ovules. These
ovules both shared a common outer integument, but each had its own inner
integument.
The aril develops from the funiculus/funicular end of the ovule (Soegeng-
Reksodihardjo 1962; Soepadmo and Eow 1977). In D. zibethinus and the
majority of wild species, the aril eventually completely surrounds the seed; D.
griffithii, D. oblongus and D. malaccensis have incomplete arils, and D.
singaporensis has no aril (Kostermans 1958b).
The major storage reserves of durian seeds are starch and protein. No
lipid bodies are present in the storage parenchyma cells of mature durian
seeds (Brown 1995b).
Leaves : Durian leaves are oval-oblong in shape with an acuminate tip. The
leaves of D. zibethinus are hypostomatic (Shanmukha Rao and Ramayya 1981).
The stipules of durian leaves are sub-falcate and deciduous (Davis and
Bhattacharya 1974). Durian leaves have several rather noteworthy characteristics.
Rumphius (1741) noted that durian leaves have distinctly swollen petiole
bases, the anatomy of which has been investigated by Funke (1931). The
vascular system of durian petioles is very complex (Solereder 1908).
Durio — A Bibliographic Review 31
Clear spots occur on the upper and lower surface of durian leaves.
Bottle shaped mucilaginous cells in the upper and lower epidermis of durian
leaves are responsible for this phenomenon (Radlkofer 1886, 1890; Solereder,
1908). Only the narrow neck of these cells lies in the epidermal layer, the
bulbous inner portion lies in the hypodermis (Radlkofer 1886). Radlkofer
(1886) reported similarly constructed clear spots in the leaves of Boschia
(now Durio).
Lamarck (1786) stated that the bottoms of the leaves and young
stems are covered by scales similar to those of Capparis. This, among other
things, led him to ally Durio with the Capparaceae. Similar parallels were
echoed by Radlkofer (1884) because of similarities in flower construction,
scales, and leaf folding between Durio and Capparis. A brief description of
the peltate scales ‘schildharre’ of durian was first given by Bachmann (1886).
Schumann (1895) described peltate scales ‘schuppe’ in Durio, and stellate
hairs in Adansonia. He reported that both forms could be found in the tribe
Matisieae. Some typical Malvaceous and Bombacaceous hairs are described
and depicted by Solereder (1908). Salma (1991) recorded five different types
of trichome morphology on the leaves of Durio, which can be useful in the
identification of species; unfortunately, no key was given. The adaxial (upper)
surface of durian leaves are smooth and apparently hairless.
Tree architecture : Durian trees are large forest trees, which can reach heights
of 37 m (Foxworthy 1927), the first branch can be as much as 18-21 m off
the ground (Foxworthy 1927). The trunks of most species are normally buttressed
in mature specimens. Out of over 300 trees measured, an average diameter of
56 cm and a maximum diameter of 107 cm was recorded (Foxworthy 1927).
Larger specimens of Durio often become buttressed (Soegeng-Reksodihardjo
1962). Many Bombacaceous trees are classified as having the architectural
model of Massart. D. zibethinus has been assigned to the model of Roux
(Hallé et al. 1978), as has D. griffithii (Ashton 1978). The model of Roux is
characterized by having a monopodial orthotropic trunk and plagiotropic lateral
branches. Roux’s model differs from that of Massart largely due to diffuse
growth rather than rhythmic growth. Detailed description of the architecture
of D. zibethinus with diagrams is presented by Buisson (1986). The orthotropic
axis of durian seedlings is fast growing, producing numerous strongly plagiotropic
laterals; some orthotropic laterals are also produced which eventually compete
with the main axis (Subhadrabandhu et al. 1991).
of 2n=28 for durian. The authors attribute this difference to the sample of
Mangenot and Mangenot (1958) being tetraploid. As neither groups of authors
mentions the clone (if any) from which sample material was obtained, it may
be that different durian clones have different levels of ploidy. To the best of
my knowledge, this has never been investigated, however, it is certainly worthy
of further investigation. Soepadmo (1979) reported a chromosome number of
56 in D. zibethinus and 60 in D. griffithii. Cullenia excelsa Wight [=Durio
ceylanicus Gardn.] has a chromosome count of n=28 (Pushparajan et al. 1986),
thus there may be variation in chromosome numbers between the species.
This question never appears to have been directly addressed, but could possibly
be of value in the classification of the wild species of Durio.
Edibility : Of all the known species of durian, the six commonly listed as
producing edible fruit are D. dulcis, D. grandiflorus, D. graveolens, D. kutejensis,
D. oxleyanus and D. zibethinus (Soegeng-Reksodihardjo 1962). Several other
lesser known species also bear edible fruit, such as D. testudinarum and D.
lowianus (Kostermans 1958b; Ogata 1978). Fruits of the newly discovered D.
macrantha are also edible (Anon. 1992; Kostermans 1992a,b). In addition to
these species, the fruits of D. excelsus (Korth.) Bakh. and D. pinangianus
Durio — A Bibliographic Review 35
(Becc.) Ridl. may be edible (Kunkal 1984), as are the seeds of D. carinatus
Mast., the arils of which are inedible. Kostermans (1958b) mentions that the
bright red arils of D. excelsus (Korth.) Bakh. and the pink arils of D.
pinangianus are tasteless, and the yellow arils of D. lanceolatus are almost
tasteless. The fruits of D. wyatt-smithii Kosterm. have been incompletely
described. It is possible that they are also edible as this species is very closely
allied with D. zibethinus. Beccari (1889, 1921) described his discovery of a
single tree of what he termed D. carinatus var. bintulensis Becc., which had
edible fruit (D. carinatus itself has inedible fruit) (Endert 1927a). Kostermans
has apparently made no comment on the nature or affinity of this specimen
and, according to Soegeng-Reksodihardjo (1962) since Beccari’s description,
no tree has ever been found. It is possible that other recognized species have
edible fruit as the fruit is completely unknown, or has never been described
for D. burmanicus, D. crassipes, D. kinabaluensis, D. macrolepis and D.
purpureus. Additionally, the mature fruit has never been collected or described
for several other species.
The amount of mass that the aril contributes to the entire fruit (i.e.,
the amount of edible portion of a durian fruit) varies greatly with the clone
or variety of fruit. Some recorded values are 30% (Adriano 1925), 19% (Abdullah
and Ragab 1970), 29% (Joachim and Pandittesekere 1943), 30% (Pratt and
Del Rosario 1913) and 14.5% (Intengan et al. 1955).
Apart from being eaten fresh when ripe, the aril is also traditionally
mixed with coconut juice, sugar, rice flour and eggs to make a cake like
concoction (dodol) (for recipes, see Soegeng-Reksodihardjo 1962; Coronel et
al., 1983; Anon. 1986b). Durian cake is now a popular contemporary commercial
product (Paweenakarn et al. 1992). Nutritional information for durian cake
has also been published (Leung et al. 1972; Ismail and Seow 1982; Paweenakarn
et al., 1992; Seow 1994). Several other commercial durian products are produced
Durio — A Bibliographic Review 36
including candy and jam. The author has also encountered durian filled swiss
rolls and donuts. At the present time, durians are not specifically grown for
the production of processed products (Maneepun et al. 1994), most commercial
durian products are likely produced using lower grade, damaged or surplus
fruits. Unripe arils are apparently eaten after roasting by the Bataks of Sumatra
(Ochse and Bakhuizen Van Den Brink 1977). Unripe arils are also recorded
as being used as an ingredient in soup (Ochse 1961). The ripe arils can be
prepared by fermenting them inside bamboo-joints, or earthenware vessels
usually for 3 to 4 days, but in some locations up to 2 weeks (Ochse and
Bakhuizen Van Den Brink 1977). The bamboo joints are either buried during
this time, or suspended over a source of smoke. The following comment from
Burkill (1966) would seem to adequately summarize the previous recipe:
‘Apologists say that the fruit should be eaten before the garlic flavour is at all
apparent. It must be added that to the depraved taste of the Besisi fermented
durian pulp, obtained by burying the aril in a time of glut, appeals’. Wallace
(1856, 1869) stated that durians were sometimes preserved with salt in jars or
bamboo joints by the Dyaks of Borneo. In the Moluccas, fish is sometimes
flavoured by smoking it above empty durian husks (Ochse and Bakhuizen
Van Den Brink 1977).
Nitrogen 0.211%10
(Contd....)
Durio — A Bibliographic Review 38
Table 3. Contd.
Minerals 1.2%5
(Contd....)
Durio — A Bibliographic Review 39
Table 3. Contd.
Chlorine 4 mg/100g23
K2O 70 mg/100g23*
(Contd....)
Durio — A Bibliographic Review 40
Table 3. Contd.
The edible aril of the common durian (D. zibethinus) is a good source
of vitamin C (Table 3), which is of the order of 33 mg per 100 g of aril
(Abdullah and Ragab 1970). This is roughly equivalent to the vitamin C
content of many citrus fruits. Phang (1976) estimated that durian arils contain
100 mg of vitamin C per 100 g of aril, however, he also warned that this
estimate was likely inflated due to interference from mercaptans, naturally
present in the aril, with his assay. This is likely a problem associated with all
published estimates of the vitamin C content of durians, thus all published
values are likely overestimates. Rosedale (1935) used a Guineapig bioassay to
estimate the vitamin C content of durian and various other foodstuffs. He
assigned an antiascorbic value to durian of 10 grams (the amount of durian
necessary to alleviate the symptoms of scurvy) which was approximately only
one third that of citrus fruits. This result supports the suggestion that published
vitamin C contents for durian are overestimations.
Fresh durian arils contain 2-2.5% protein (Table 3). The total and
essential amino acid composition of durian arils has also been investigated
(Zanariah and Noor Rehan 1987) on a ‘per gram fresh weight’ basis. The data
show that durian is a better source of all the essential amino acids than dates,
peaches, oranges, mangoes, cempadak or papaya (Table 4). However, it should
be remembered that durian has a much higher percentage of total protein than
all these fruits.
Durio — A Bibliographic Review 42
The fruit has been examined and tested positive for thermostable
anti-thiamine (thiaminase) activity (Rattanapanone 1979). Bate-Smith (1959)
described the presence of caffeic acid and small amounts of leuco-anthocyanins
in durians (although this was probably from an analysis of the leaves). Baldry
et al. (1972) listed several alcohols present in the aril including ethanol,
methanol and n-propanol. Wong and Tie (1995) have recently questioned the
presence of ethanol and methanol. Numerous other compounds have been
identified in fresh durians (Table 5). The creamy consistency of the aril is
attributed to gums, pectins and hemicelluloses (Martin 1980).
(Contd....)
Durio — A Bibliographic Review 44
Table 5. Contd.
(Contd....)
Durio — A Bibliographic Review 45
Table 5. Contd.
Propyl 2-methylbutanoate
(N-propyl a-methylbutyrate) 1,3
Propyl 2-methylpropanate 3
Propyl propanoate
(N-propyl propanoate) 1,3
S-propyl thioacetate 3
S-propyl thiopropionate 3
2,4,6-trimethyl-1,3,5-trithiane 3
1=Baldry et al. (1972), 2=Moser et al. (1980), 3=Wong and Tie (1995).
sinapic acids are reported absent and leuco-anthocyanins present in low amounts
or absent (Bate-Smith 1959).
Fatty acids : The fresh arils of durian fruits consist of about 3-5% fat (Table 3).
The exact amount of fat seems to vary between clones as does the nature of the fats
themselves. The aril of the durian was first analyzed using gas chromatography by
Aspiras and Tocino (1971), but no conclusions about its constituents were made.
The doctoral thesis of Greve (1974) included an analysis of the fatty acid
composition of durian seeds, arils and husks collected from two locations in
Thailand (Table 6). Not only does the fatty acid makeup differ between these three
tissues, but it differed markedly between the two varieties of durian. Most notably,
20:0 fatty acids constitute approximately 33% of the total fatty acids found in the
husk (pericarp) and almost 10% of that of the seeds in Prajeen Rayong durians, but
were found in negligible amounts in all tissues of Chanthaburi durians (Greve
1974); variance in the fatty acid makeup of the arils was much less between these
two varieties than that found in the pericarp and seeds. Berry (1980a,b,c) reported
on the lipid constituents of both the arils and seeds of durian, in particular the
presence of cyclopropene fatty acids which are present in many related genera,
and which may be carcinogenic (Berry 1980b). He concluded that cyclopropene
fatty acids are present in the seeds in unusually high quantities, 65% of the total
fatty acids in durian seed oil are cyclopropenoids (Berry 1980a). Cyclopropenoid
fatty acid content was reduced, but not eliminated upon cooking. Sterculic,
dihydrosterculic and malvalic acids were present in the uncooked seeds but not in
the aril. Due to the toxic and perhaps carcinogenic nature of these substances,
Berry (1980b) concludes that it would be unwise to ingest uncooked durian seeds.
Berry (1981) compared fatty acid composition of four durian clones and linked the
palmitic:palmitoleic acids ratios with taste. The lower this ratio, the higher fruits
were ranked by a panel of tasters. He also noted that the degree of saturation of
fatty acids varies between clones, which may influence retention of volatile
flavouring compounds, and hence the flavour itself. However, at present, this
relationship remains correlative rather than causative.
Durio — A Bibliographic Review 46
Smell : Although the fruits of Durio zibethinus are infamous for their foul
smell, they apparently are not the most offensive of all the known species.
Most species of Durio produce odourless, or nearly odourless fruits, however,
the odour of the ripe (and edible) fruits of Durio dulcis is apparently so vile
and pervasive that they can be smelled for miles (Soegeng-Reksodihardjo
1962); Kostermans (1958b) describes the smell of these fruits as ‘simply
nauseating’. Hayes (1957) recorded that it is believed that the smell of durian
fruits can be eliminated by soaking the flesh in coconut milk overnight. I
have, however, seen no recorded experiments on this procedure.
Stanton (1970) and Baldry et al. (1972) divided the smell into two
distinct categories, a very strong onion-like smell (alliaceous), probably caused
by thioethers; and a second fruity smell due to the presence of esters. Stanton
(1966) suggested that indoles and skatols may be present in the fruit accounting,
in part, for its malodorous nature. Stanton (1970) presented evidence that the
arils do not release thiols, and thus thiols are not responsible for the smell.
However, esters and thioethers were implicated. Baldry et al. (1972) analyzed
durians obtained from Singapore which gave similar results, but a durian
obtained from Kuala Lumpur was found to owe a large portion of its odour
to thiols with minimal contribution by thioethers. The major volatile components
were found to be ethanol, n-propanol and ethyl a-methylbutyrate [=ethyl 2-
methylbutanoate]. This issue was further addressed by Moser et al. (1980)
who provided evidence that durian arils do not contain thiols and that mature
arils do release hydrogen sulphide, while those of immature durians do not.
They also provided evidence that the release of hydrogen sulphide is not due
to bacterial action in the fruit (as has occasionally been suggested), since
tissue from immature arils did not yield any colonies when cultured on substrates
for H2S forming species. Martin (1978) stated that hydrogen sulphide and
diethyldisulphide are responsible for the foetid odour. Durians smell more as
they mature; analysis of H2S production by 500 grams of durian at different
stages of maturity produced the following results: immature durian arils produce
0.009 g H2S, ripe durian arils 0.023 g and very ripe durian arils 0.047 g
(Greve 1974).
Indoles were not detected as was predicted by Stanton (1966). The polysulphides
identified by Moser et al. (1980) were mainly ethyl polysulphides which
apparently rarely occur in fruits. Unfortunately, Moser et al. (1980) obtained
their durians from Thailand. Thai durians differ substantially in odour and
flavour from Malay durians, the Thai durian being sweeter and considered
less aromatic than Malay clones (Mohamad Idris 1987). Thus, the differences
in constituents found between the study of Moser et al. (1980) and Baldry et
al. (1972) may either reflect clonal differences in aroma composition or
differences due to technique.
Wong and Tie (1995) were unable to confirm the presence of several
of the volatile compounds reported in both previous studies including ethanol,
methanol, ethyl methacrylate and several sulphur containing compounds. They
attribute these differences to better technique, and to using ripe durians and
a minimal time between harvest and analysis. There are three interesting
aspects of this study, asides from a long list of new volatile components.
Firstly, Wong and Tie (1995) sampled and compared the composition of
fruits from three named clones, D15, D28 and D74. Not only did they show
considerable differences in the total yield of volatile compounds between
clones, but numerous large differences in quantities of particular volatiles
exist between these clones. Some volatiles were not detectable in all of the
clones. Secondly, Wong and Tie (1995) demonstrated the presence of three
different a-hydroxyketones which they claim have never before been reported
from an analysis of a fruit. Thirdly, these authors stated that no trace of
H 2S was detectable in the aroma profile of the fruits from any of the three
clones tested. This result is at variance with several previous works (Baldry
et al. 1972; Greve 1974; Martin 1978; Moser et al. 1980) and is likely to
be related to change in aroma composition over time during ripening of the
fruit. The H2S production increases markedly as the fruit ripens (Greve 1974).
Thus, although Wong and Tie (1995) used ripe fruit, it may not have been
ripe enough to produce H2S. Apart from the study of Greve (1974), the change
in aroma composition of durian fruits over time has not been examined. Such
Durio — A Bibliographic Review
Table 6. Fatty acid composition of durian
Tissue Clone Saturated Palmitic 14:0 15:0 16:0 16:1 17:0 18:0 18:1 18:2 18:3 20:0 22:0 Dihy- Mal- Ster- Cis
unsatur- Palmit- drost- valic culic vaccenic
ated oletic erculic
Aril D241 1.4 4.7 0.5 <0.1 39.8 8.5 <0.1 0.8 45.8 1.8 2.7
Aril D2 1 1.7 6.9 0.5 <0.1 35.9 5.2 <0.1 1.0 51.0 2.4 4.1
1
Aril D66 1.8 5.3 0.8 <0.1 33.8 6.4 <0.1 0.9 48.7 2.8 7.0
Aril D8 1 1.9 13.9 0.5 <0.1 32.3 2.3 <0.1 2.2 53.6 3.2 6.0
Aril Unknown 0.91 34.13 7.1 1.21 42.14 7.85 5.69 <0.01
12
Aril Unknown 0.34 28.94 5.16 1.23 5.98 3.16 2.21 <0.01
22
Whole Unknown2 82% 0.12 12.20 1.15 1.42 8.42 6.50 11.3 1.21 2.52 15.72 38.53
fresh unsaturated
seeds
Pericarp Unknown 30.9 4.2 1.8 52.8 4.2 4.0 <1
Thai#13
Pericarp Unknown 14.9 1.4 5.1 17.8 7.6 4.9 33.2
Thai#23
Aril Unknown 33.9 4.9 1.0 51.2 3.0 5.1 <1
Thai#13
Aril Unknwon 31.9 6.9 1.0 51.0 3.0 5.4 <1
Thai#23
Seed Unknown 26.8 8.4 3.3 38.8 5.9 3.0 <1
Thai#13
48
(Contd...)
Table 6. Contd.
Seed unknown 23.0 3.1 1.2 41.0 11.5 5.3 9.5
Thai#23
Aril unknown 0.6 31.3 3.4 2.1 (47.1, 1.3 7.1 8.4
Thai4 4.3)*
Fatty acid compositon of various parts of durian fruits. Note: all values are in percent GLC peak area except for that of Shibahara et al. which is
in % wt from GC analysis.
1
Berry (1980c) and Berry (1981).
2
Berry (1980b) (Unknown#1 was from a Malay durian with yellow arils, #2 was from a Malay durian with cream coloured arils).
3
Moser et al. (1980) (Thai#1 was a durian from Chandburi, Thai#2 was from Prajeen Rayong.) Note: This data is originally from the thesis of Greve
(1974).
Durio — A Bibliographic Review
4
Shibahara et al. (1987) *Note: Vaules are for 18:1 n-9, n-7 respectively.
49
Durio — A Bibliographic Review 50
Miscellaneous uses of fruit : Durian husk extracts have been studied for
their suitability as aqueous binders for granules and tablets (Umprayn et al.
1990a,b). These authors showed that husk (pericarp) extracts are suitable
for the preparation of granules and to have desirous binding properties for
the manufacture of tablets (Umprayn et al. 1990b). The ashes of D. zibethinus
are used to extract a dye which, in turn, is used to prepare batik dye
(Kostermans 1958b). The ashes have also been used for whitening silk
(Kostermans 1958b; Morton 1987). The empty husks are also used in Java
as a source of fuel (Ochse and Bakhuizen Van Den Brink 1977; Morton
1987).
Durio — A Bibliographic Review 51
More recently, a Thai research paper has again studied the effect of
alcohol-durian mixtures on mice and rats (Nilvises and Saengsirinavin 1986).
Similar results to those of the previous studies were found. Furthermore, this
study revealed that similar results are obtained with durian fruits of several
different clones.
Vermifugal properties : Hurrier and Perrot (1907), and Morton (1987) both
report that durian is used as a vermifuge. De Padua et al. (1978) reported it
to be a vermifuge, vermicide and an anthelmintic. I have, however, been
unable to locate any prescription for deworming based on durians, or to find
any record of what part of the plant is supposed to have this property.
Treatment of jaundice : It appears that the leaves have been used in the
traditional treatment of jaundice (Brown 1954). Burkill and Haniff (1930)
have recorded the following traditional prescription for the treatment of jaundice:
‘Boil leaves in water and bathe in it.’
Thus, they are not optimal food items for persons with non-insulin dependant
diabetes mellitus (Roongpisuthipong et al. 1991). The reason for the greater
effect of durian on insulin levels in the blood compared to other fruits is still
unknown.
that makes one short of breath (Rumphius 1741; Kostermans 1958b; Stanton
1966). Seeds contain sterculic acid which may be responsible for this effect
(Berry, 1980b).
Seeds
Mature seed constituents : The moisture content of fresh durian seeds is
enumerated by Chin et al. (1989). The moisture content of the embryo is
given as 65% (w/w) on a fresh weight basis, while the value for the ‘whole
seed’ is only 50%. The authors do not define what is meant by whole seed,
so it is unclear if this refers to the embryo and seed coat, or the embryo,
seed coat, and the large fleshy aril. Berry (1980b) gave measurements of
durian seed moisture as 77%, Soepadmo and Eow (1977) stated that the
average moisture content of fresh seeds is 51%, Chin et al. (1984) listed the
moisture content of freshly harvested seeds as 39%. The large variance in
these estimates is likely due to differences in drying regimes. Estimates of
seed moisture content of durian apparently vary greatly when the drying
time and temperature is varied (Grabe 1992). Fresh durian seeds contain
some small amounts of protein, 1.57% (Berry 1980b), but consist largely of
starch (Table 7). Mature seeds appear to be metabolically active when shed;
polysomes are evident in mature seeds and specific enzymatic activity is
detectable (Brown 1995b).
Culture of seeds and seed size : The seeds of durian are large. According to
Troup (1921), about 1 dozen seeds weighs 454 g. I think this may be an
overestimation unless the author made his calculations from a variety with
unusually large seeds. From my own measurements, mature durian seeds weigh
approximately 20 grams. Davis and Bhattacharya (1974) give an average
weight of 12 to 15 grams from measurements made on the seeds of two
fruits4. Chin et al. (1984) state that the weight of 1000 durian seeds is 14783
g (14.7 g per seed). There is a great variation in seed size within a single
4
These estimates include the smaller semi-aborted and full sized seeds.
Durio — A Bibliographic Review 57
durian fruit, some seeds abort at different stages of development. Full sized,
but flattened and completely empty seed coats may often be found inside the
arils of mature fruits. Fully mature and filled durian seeds sink when immersed
in water, but ‘aborted’ or ‘partly-filled’ seeds float (personal observation).
Partly filled seeds can often be successfully germinated (personal observation).
When mature, the seeds are non-endospermous, the endosperm becoming
exhausted several weeks before fruit abscision. The endosperm remains in a
free nuclear state until late in embryogeny at which time it becomes cellular
(Soepadmo and Eow 1977). According to Ho (1972), there is a great deal of
genetic variability in endosperm abortion in Durio, however, I think that he
is actually referring to seed abortion. The seeds of durian are described and
portrayed diagrammatically by Corner (1976) with particular attention to the
vascular supply, in his encyclopedic ‘Seeds of Dicotyledons’.
susceptible to oxidative browning, thus, excised embryos fared best when first
soaked in antioxidant, and subsequently grown on charcoal containing media,
containing 1.0 mgl-1 NAA (a-napthalene acetic acid) or IAA (indole-3-acetic
acid) and 1.0 mgl-1 kinetin, BAP (benzylaminopurine), or 2iP (2-isopentyladenine)
(Chin et al. 1988). Embryos of D. lowianus cultured on woody plant medium
supplemented with BAP produced callus (Kongnakorn et al. 1985). Apparently,
some success in callus formation from excised cotyledonary tissue of D. zibethinus
has also been achieved (Rao et al. 1982; Johri and Rao 1984), however, as of
yet no literature has been uncovered on plant regeneration from such callus
cultures of durian5.
Viability : It has long been known that the seeds of durian have a short period
of viability (MacMillan 1909; Main 1909a,b; Troup 1921; Chevalier 1934).
The seeds of durian are now categorized as recalcitrant (Hofmann and Steiner
1989); seeds with a relatively high moisture content at maturity, which cannot
withstand desiccation (Hanson 1984; Roberts et al. 1984), and thus have a
relatively short period of viability. In durian, this is correlated with the inability
to withstand chilling or freezing. The recalcitrant nature of durian seeds presents
major problems in the storage of durian seeds both for commercial purposes,
and for conservation. As durian is undoubtedly suffering some degree of
genetic erosion (Sastrapradja 1975), research into the physiological response
of durian seeds to chilling and drying is assuming a new importance.
Seeds stored at 36oC lose viability after 6 days, while surface sterilized
seeds sealed in an airtight container under nitrogen can maintain viability for
32 days if the temperature is lowered to 20oC (Soepadmo and Eow 1977).
Further experiments by Teng (1980) showed that fresh durian seeds, stored
for 3 months on wet tissue paper at 15oC, maintained 79% germinability,
although some problems with fungal infection and radicle protrusion were
encountered. Storage of durian seeds at 5oC for 10 days resulted in a reduction
of viability to 80%. Viability was completely lost at this temperature after 20
days of storage. Seeds stored at 15oC and 29oC maintained their viability for
20 days; however, seeds stored at the higher temperature had all germinated
after 10 days. Hanson (1981) performed some preliminary drying and storage
experiments on some tropical recalcitrant seeds including durian. Durian was
shown to have a viability period at 27oC of 3-4 weeks; seeds stored at 15oC
exhibited only 15% germination after 2 months.
5
Numerous attempts at tissue culturing durian seeds have been attempted by myself and Miss
L. Sreedhar, in the Department of Botany, University of Guelph. Although vigorously growing
callus is somewhat easy to produce, various combinations of plant growth regulators at different
concentrations have proven unsuccessful in regenerating plants.
Durio — A Bibliographic Review 59
The moisture content at viability loss after slow and fast drying was
reported as 71 and 63% respectively (Boyce 1989; Grabe 1989). These values
are much higher than numerous other accounts of the minimal moisture content
of viable durian seeds. The differences between this report and those of others
are undoubtedly related to the exact method of moisture determination which
is known to greatly affect moisture determination in durian seeds (Grabe
1992).
It has been stated that the cotyledons are shed shortly after germination,
Durio — A Bibliographic Review 61
Pollination Biology
Anthesis : Durian flowers open in mid afternoon (the flowers of some clones
are reported to open as early as 2:30 PM (Tidbury 1976). Even though the
flowers open, the anthers do not dehisce and release pollen until about 7:00
PM. Polprasid (1969) states that durian pollen grains function maximally at
9:30 PM. By early morning, the calyx, corolla and staminal groups have
abscinded leaving just the pistil attached to the receptacle. Jamil (1966) reported
that the timing of anthesis and anther dehiscence varies by several hours
between clones, but no data were presented. Attempts to pollinate durian
flowers with pollen released by crushing from undehisced anthers resultd in
no fruit set (personal observations). Although pollen is not functional before
anther dehiscence, pollen does remain viable up to two days after anthesis
when stored in a refrigerator (Coronel 1966).
Valmayor et al. (1965) and Coronel (1966) reported that the stigmas
of durian are receptive to pollen 36-48 hours before the flowers fully open.
Coronel (1966) reports that hand pollination of 36-48 hours before anthesis
flowers by surgical removal of the outer floral parts leaving just the pistil,
results in a greater percent fruit set than hand pollination of fully mature
flowers. Razak et al. (1992) have shown 87.5% receptivity of D24 durian
stigmas 10 hours before anther dehiscence with 42.1% receptivity remaining
24 hours after anthesis (receptivity was estimated by measuring percentage
fruit set). Salakpetch et al. (1992) found that stigma receptivity in some Thai
cultivars began 24 hours pre-anthesis, receptivity remaining high until noon
the day after anthesis, after which it rapidly declined. Shaari et al. (1985)
reported that receptivity was reduced to approximately 50% by noon the day
after anthesis. Jamil (1966) has suggested that the period of stigma receptivity
varies considerably between clones. Salakpetch et al. (1992) showed that
pollen viability also varies between clones in a study with 4 Thai cultivars,
viability ranged from 77 to 93% two days after anthesis.
Natural pollinators : Today, durian trees are generally regarded as bat pollinated,
although Beccari (1889) was of the opinion that birds could be pollinators.
Durio — A Bibliographic Review 62
The first suggestion of bat pollination of Durio is that of Beccari in his book
‘Malesia’ (1889) who records Macroglossus minimus Geoffr. visiting the flowers.
This account apparently did not make a big impression for it was not until
over 40 years later, that Boedijn and Danser (1929) noted a flock of bats
visiting a durian tree, their activity causing the shedding of numerous stamens
and calyxes. Although they didn’t observe or demonstrate actual pollination
of flowers by bats, they postulated that this may have been the case. Interestingly
enough, the affinity of flying foxes for the nectar of durian flowers must have
been common knowledge, an account of their feeding is given in an illustrated
tour guide to the Federated Malay States first published in 1910 (Harrison
1910a). Van Der Pijl (1936) conclusively demonstrated that durian flowers
have typical bat pollinated flower morphology, and that the visitation of flowers
by bats does indeed cause fruit set; he also raised strong arguments against
birds as possible pollinators. As birds have not been shown to visit durian
flowers during the time when pollen has been released, they are unlikely
natural pollinating agents.
Self-incompatibility : It has been known for some time that some durian
trees are self-incompatible (Coronel 1966); although crossing and selfing
tests with named clones have been conducted (Jamil 1965), no details
seem to have been published. This is unfortunate as such information
would be useful in determining good clonal mixtures for orchards and for
the production of better clonal material in the future. The Malaysian clone
D99 has been shown to be self-compatible (Zainal Abidin and Nik Masdek
1992). Clone D24 (Shaari et al. 1985), and the Thai cultivars Gumpun and
Luang (Lim el al. 1992) have been demonstrated to be self-incompatible.
However, compatibility information for the many other clones/cultivars is
not available.
Heterostyly : Chin and Phoon (1982) claimed that there is heterostyly between
different durian clones, although they presented no evidence for this statement.
My personal observations (unpublished) of the stylar lengths of different durian
clones show a lack of distinct size groupings between clones. Lye (1980)
described several differences in stylar morphology that apparently exist between
clones, but he did not mention stylar length. The study of Shaari et al. (1985),
as mentioned previously, showed that unfertilized ovaries abscinded much
earlier than did those of selfed flowers of a self-incompatible clone. Heterostyly
would not explain this observation. Personal observations, and the work of
Lye (1980) have shown that there is definite heteromorphy in stigma and
styles between clones. Despite these differences in stylar morphology, selfed
Durio — A Bibliographic Review 65
pollen has been shown to germinate on the stigmas of selfed flowers (Namuco
1978). Furthermore, pollen grains collected from fallen staminal phalanges
will germinate after storage at room temperature for 48 hours, thus stigmatic
exudate is not strictly necessary for germination. Thus, it seems unlikely that
stigmatic heteromorphy contributes to or is associated with self-incompatibility
mechanisms.
Premature fruit drop : As mentioned above, premature fruit drop occurs after
all abscissions due to a lack of fertilization or self-incompatibility. Yaacob et
al. (1978) suggested that it may be due to some physiological disorder within
the aborted fruits; however, pathology may not be necessary to explain this
phenomenon. The number of harvestable fruits is not necessarily correlated with
the number of successful pollinations above some basal level. As previously
mentioned, the fruit set of durian flowers is quite low (Soepadmo and Eow 1977;
Mardan and Zainal 1986). Other authors have reported very high percentage
fruit set using artificial pollination (Valmayor et al. 1965; Coronel 1966). The
discrepancy between these high values and the low values previously described
Durio — A Bibliographic Review 67
probably lies in the number and distribution of flowers pollinated. Durian trees
produce large number of flowers, but owing to the large size of a mature durian
fruit, it is undoubtedly impossible for all flowers produced to develop into mature
fruits, even if the fruits are all healthy. Polprasid (1969) noted that trees with
more flowers had a lower percentage fruit set. Thus, even if all flowers on a
tree are suitably pollinated, the tree must, and apparently does, have some
mechanism to selectively abort some of its excess fruits. The study of Chandraparnik
et al. (1992) showed that trees treated with paclobutrazol and thiourea had a
more even distribution of flowers throughout the canopy, and had a larger
number of developing fruits at 5 weeks after pollination as compared to control
trees.
Leaf flushing : Durian trees typically produce a flush of new leaves 3-8
weeks after anthesis (Salakpetch et al. 1992). Leaf flushing in durian is
thought to contribute to premature fruit abscision and hence decrease percentage
fruit set. To test the validity of this assumption, and the capacity of plant
growth regulators to delay the onset of leaf flushing, foliar sprays of several
substances were tested on mature durian trees (Punnachit et al. 1992). 2500
ppm mepiquat chloride (300 g/20 l), KNO3 and 2500 ppm daminozide, and
(500 g/20 l) low nitrogen (0-52-34) foliar fertilizer were found to be effective
Durio — A Bibliographic Review 68
Species which feed upon durians : Some researchers have provided information
regarding the species of animals which naturally feed upon durian fruits.
Hubback (1941) reported that elephants trample the fruits and then eat the
orangutans fed on durians (Wallace 1869; Davenport 1967; Shetford 1985).
Rijksen (1978) records that orangutans also eat the fruit of Durio oxleyanus,
which constitutes an ‘esteemed’ food item of the orangutans of Ketambe.
Orangutans have developed behaviours to aid in the plucking, carrying and
opening of durian fruits (Rijksen 1978). Corner (1964) enumerates rather
graphically several species of animals which feed upon durian. According to
Barrett (1912), wild pigs eat durian fruits. A species of Philippine flying
squirrel (Sciuropterus mindanensis) is known to feed on the fruits of D. zibethinus
(Rabor 1939). Gardner (1847) recorded that monkeys eat the seeds of Durio
ceylanicus Gardn. [=Cullenia ceylanica (Gardn.) K. Schum. sensu Kostermans].
Durio — A Bibliographic Review 69
Kostermans (1953b) states that most of the fruits of wild species of durians
are lost by damage from squirrels, monkeys, tupais and hornbills. Hawkins
(1986) claims that durians are eaten by elephants, tigers, deer, rhinoceros and
monkeys. Watson (1984) claims the fruit is even attractive to the domestic
cat.
Natural dispersal of seeds : Ridley (1894) speculated on the dispersal of
several durian species by bears, birds (especially hornbills) and even turtles.
D. zibethinus is perhaps dispersed by bears. The wild bear, Helarctos malayanus,
is known not only to eat fallen fruits, but even to climb trees to obtain durians
(Ridley 1894). Ridley further speculated that D. oblongus seeds are dispersed
by birds, as it is native to Singapore where there are no bears. Ridley related
that upon feeding a fruit of this species to a wild bear (Helarctos malayanus),
it ate the arils with great gusto but refused to eat the seeds. An unnamed
Durio species [Ridley described it as having small red arils=D. graveolens?]
is probably dispersed by hornbills. This sentiment is echoed by Browne (1955):
‘I well remember being seen out of a durian grove by a tiger.’ Whitmore
(1990) said: ‘Tigers are notorious for their passion for durians’. Rutten (1939)
reported the germination of durian seeds in elephant faeces, but none became
established.
Meijer (1968) noted that the fruits of the cauliflorous D. testudinarum,
which bears fruits at the base of the tree, are widely believed by natives to be
eaten by tortoises. Ridley (1894) recorded that this species is known in Borneo
as the tortoise durian.
The seeds of D. oxleyanus are severely damaged by orangutans in
attempts to open the fruit (Rijksen 1978). Rijksen (1978) suggests that the
sun-bear and tigers are probably the main dispersers of seeds of this species.
Fruiting Seasons
Maturation of buds : The floral buds take up to 5 months to become mature
fruits. In my experience, mature fruits of clones D8 and D24 require 95-110
DAP (days after pollination) to mature. D99 requires only 95-100 days (Zainal
Abidin and Nik Masdek 1992). Mon Thong durians are harvested 120-130
DAP (Chattavongsin and Siriphanich 1990a). Durians typically have two seasons
during the year, a major and a minor one. The same trees do not necessarily
bear fruits in both seasons (Browne 1955).
Several authors have recorded, for various locations, the months in
which durian trees typically flower and fruit (Table 8). The maturation of durian
fruits is traditionally indicative of the end of the dry season (Dove 1985).
Floral buds are produced well in advance of flowering and are usually
dormant for at least one year, the breaking of bud dormancy is facilitated by
Durio — A Bibliographic Review 70
cool nights (Ong and Lee 1981). Some floral buds do, however, develop from
initiation to anthesis without interruption by a dormant state (Subhadrabandhu
et al. 1991). Occasionally, floral initials can revert back to vegetative growth
(Subhadrabandhu et al. 1991).
D. griffithii has synchronized flowering, although some individuals
produced small numbers of flowers asynchronously (Yap 1980).
Table 8. Durian seasons in different geographic areas
Malaysia
Sabah Fruiting August to December, peak in October (Davenport
1967)
Sarawak Main fruiting season January-March (Mohamad Idris 1987)
Peninsular Main fruiting season June-August (Mohamad Idris 1987)
Peninsular Two fruiting seasons June-July, August (Anon. 1982b)
Penang Fruiting June-August (Low 1836)
Jambu Rias, Pahang Flowering in April (major) and August (minor), fruiting
August and February. The major fruiting season harvest
in one year was 266.6 kg/hectare in February and 2269
kg/hectare in August (Chiow 1976).
Philippines
Mindanao and Sulu Flowering May-June, fruiting August-November (Rodrigo
1968; Galang 1955)
Los Banos Flowering February, fruiting July to August (Namuco 1988)
Davao City Flowering January to May, fruiting May-September (Pascua
and Cantila 1991)
(Contd....)
Durio — A Bibliographic Review 71
Table 8. Contd.
India
Burliar Research Station Fruiting July to September (Naik 1949)
Manipulating seasonality : Durians are only available during short day periods
of the year. Attempts have been made to alter the timing of flowering to
spread out the season and, therefore, make durian fruits more readily available
throughout the year. Zainal Abidin et al. (1986) have suggested several ways
of reducing the effects of seasonality on the price and availability of durians.
They recommend exploiting differences in fruiting seasons that exist in different
regions of Malaysia, as well as some useful characteristics of some Malaysian
clones; for example, the ability of some clones to fruit twice a year and the
use of early mid and late producing clones to extend the season. Previous and
unobtainable work cited in Chandraparnik et al. (1992) has apparently shown
that ethephon, daminozide, NAA (a-napthalene acetic acid), or fertilizer
application are unsuccessful in inducing early flowering in durian; GA3
application has an adverse effect on early flowering. Chandraparnik et al.
(1992) investigated the effect of foliar sprays of paclobutrazol (Cultar ®), a
suppressor of gibberellin (GA) production, on early flowering and fruit production
on the Thai variety Chanee. They found that increasing concentrations of
paclobutrazol positively affected the number of flowers produced per tree.
Flowering of treated trees (using the highest concentration of paclobutrazol)
began up to 43 days before that of control plants. However, the number of
Durio — A Bibliographic Review 72
harvestable fruits produced, and average fruit weight was negatively affected
by this treatment, treated trees also took 2 extra weeks to mature fruits.
Thus, even at the highest concentrations of paclobutrazol used, a maximum
of one month advance on mature fruit production over untreated trees was
gained due to the extended developmental time. Chandraparnik et al. (1992)
also investigated the effects of foliar sprays of thiourea on previously
paclobutrazol treated trees. A somewhat linear relationship between the
concentration of thiourea and the number of flowers produced was shown.
Flower density was increased up to 400% over control trees, and the distribution
of inflorescences throughout the tree was found to be more even than in the
control treatment. The average number of fruits per tree at 5 weeks after
anthesis was significantly increased, however, no indication of the effects on
the numbers, size or quality of harvestable fruits was given. Lin (1992) proposed
an irrigation forcing model to delay the production of durian inflorescences.
This model has not been field tested.
Durio species are found growing in lowland and hill primary forests in
Malaysia, up to 1000 m, at a density of not more than 3-4 trees per hectare
(Soepadmo and Eow 1977). Durians (species not named) were recorded at a
density of 3.7 trees per 40 ha in Ulu Kelantan forest, Malaysia (Whitmore
1990). Foxworthy (1916) states that 1.07% of the forest trees of the East coast
of Borneo are Durio sp. based on an examination of almost 690 ha. In 1967,
durian made up approximately 1 tree per 2.6 per km2 in Sabah (Davenport
1967). The relative density of D. griffithii Mast. at Bukit Sebelah in Sumatra
is recorded at 4.16 tree/ha (Mukhtar et al. 1990). Durio sp. tree densities of
4.4, 3.0 and 2.2 trees per hectare are recorded at Pasoh Forest reserve, Taman
Negara and Krau games reserve respectively (Soepadmo 1979).
large trees per hectare while the latter was a dominant species with up to 30
large trees in 2 hectares.
Sutisna and Soeyatman (1985) examined a logged-over peat swamp
forest, five years after logging, in 3 locations in Eastern Sumatra, Indonesia;
Suakandis (Jambi Province), Sei Teban (Riau province), and Sei Lalan (South
Sumatra province). In this study, D. carinatus was found at a density of
100.0 sapling stage individuals/hectare, 9.2 pole stage/hectare, 10.3 tree
stage/hectare at Suakandis, no individuals of this species were recorded at
the other two locations. D. lowii (the authors most likely mean D. lowianus)
possibly was present only at Sei Lalan, measurements of number of individuals
per hectare are 21.43 sapling stage/hectare, 10.29 pole/hectare and 1.86 tree
stage/hectare.
Centre of diversity : Mendoza (1941) concluded that the centre of diversity
of the genus was Borneo. About 20 of the approximately 30 recognized species
are found in Borneo, 15 of which are endemic. There are 11 species which
are found in Peninsular Malaysia and 5 of it are endemic. Only two recognized
species are endemic to Myanmar. Thus, although the genus most probably
originated in Borneo, it does seem to have spread up the Malay peninsula
before all contemporary species had evolved. The centre of diversity of the
genus is not in dispute, however, opinions abound on whether D. zibethinus
is native or introduced to regions outside Borneo. Furthermore, there is some
debate as to whether this species actually exists anywhere in the wild, or
whether it is the descendant of some wild species.
Wild form of Durio zibethinus : Popeno (1920) stated that J.D. Hooker did
not think that the natural distribution of D. zibethinus extended to the Malay
peninsula, and furthermore that Hooker suspected that D. malaccensis might
be the wild form. Although both these statements appear in ‘The Flora of
British India’ by J.D. Hooker, they appear in chapter 26 on the Malvaceae
written by M.T. Masters. Regardless of who made these claims, Kostermans
(1958b) states, without explanation, that ‘Hooker’s suggestion that D. malaccensis
is the wild form of D. zibethinus is, of course, entirely wrong.’ It should be
mentioned, however, that there has been tremendous confusion as to what
exactly constitutes D. malaccensis.
Van Steenis (1949) suggested that a species he discovered in Southern
Sumatra (D. spontaneus Bakh.) is the closest wild ally of D. zibethinus.
Kostermans (1958b), however, does not recognize D. spontaneus Bakh. as a
valid species, rather he includes this specimen under D. lowianus Scort. et
King. Although this species bears fruits similar to D. zibethinus, Kostermans
does not speculate on a close affinity between the two species; rather, he
opines that his newly erected species D. wyatt-smithii Kosterm. is perhaps the
wild ancestor of the cultivated durian (Kostermans 1958b). Soegeng-
Durio — A Bibliographic Review 74
Reksodihardjo (1962) states that the ‘wild form’ of D. zibethinus need not be
one of the extant wild species.
There has also been some debate as to whether durians exist naturally
in the Philippines, or have been introduced. Wester and Barrett (1912), and
Wester (1916b, 1921) recorded D. zibethinus as confined to Mindanao and the
Sulu archipelago in the Philippines. Merrill (1926) listed Durio as a genus
represented only by introduced species in the Philippines, and Van Steenis
(1933) listed Durio as a non-native of the Philippines. Mendoza (1941)
thoroughly examined this issue, and claimed that D. zibethinus occurs naturally
on Mindanao and the Sulu archipelago as well as Palawan. Mendoza further
concluded that durian occurs only as an introduced species in the Visayas and
Luzon. Additionally, Mendoza recorded the discovery of a wild species (D.
testudinarum) growing naturally on the island of Palawan, further strengthening
his claim of endemism. Mendoza (1941) hypothesized that durians became
indigenous due to a land bridge that connected Palawan with Borneo during the
pleistocene era. Vendivil and Reynoso (1983) recently reported another wild
species, D. graveolens Becc. growing along a river in a secondary forest in
Palawan.
Durio zibethinus in the wild : Van Steenis (1949) believed that D. zibethinus has
never actually been found in the wild, and Whitmore (1990) regarded D.
zibethinus as completely unknown in the wild. Kostermans (1958b), on the other
hand, stated that D. zibethinus is probably wild in Sumatra and Borneo. This and
the question of its being native or introduced to certain regions share one property
in common, i.e., both questions are undoubtedly impossible to answer. Aboriginal
peoples have aided in the propagation of durian (knowingly or unknowingly) by
eating, trading and discarding the seeds of the fruit, probably for millennia. As
it can never be ascertained whether a tree growing in a forest grows there because
of the actions of these people or not, such arguments are purely semantic, and can
provide little on any useful information or insight.
Thailand, Southern Myanmar and a few islands in the Philippines, durians were
never introduced into New Guinea, probably because Malay and Indonesian
traders never settled in New Guinea (Knight 1980). In more recent times, durian
has been grown at the Lowlands Experimental Station in Papua New Guinea
(Bettencourt et al. 1992), and successfully grown in the Solomon Islands at the
Dala Experiment Station (8.5o South latitude) (Anon. 1968). Durian has also been
grown on the Island of Ponape in the Kolonia Botanic Gardens (7o N) (Kanehira
1935). In the Philippines, it is found mostly in Mindanao and the Sulu Archipelago
(5-10o N), but it has fruited in Laguna and Quezon provinces (Galang 1955).
Recently, durian has been introduced and successfully cultivated in Northern
Australia, however, it can only be grown in this country north of 17 o (Watson
1993).
There are two species from Myanmar of Durio, however, D. zibethinus
is probably found there only through introduction. Gamble (1881) states that it
is wild in South Tenasserim, but it is cultivated as far North as Moulmein.
According to Knight (1980), it can only grow in Tenasserim (the southern most
part of Myanmar), but it cannot grow above 16oN in Myanmar. Durian apparently
formed forests in Lower Tenasserim from 14oN southwards (Kurz 1877; Gamble
1972). Further, in the ‘Report on the Settlement Operations in the Amherst
District 1891-1892’, the durian is believed to have been introduced into the
Amherst district of Myanmar from seeds planted from a fruit washed up on the
shore from a wrecked cargo ship in the 1700s (Anon. 1893).
Durian was introduced into Britain as a greenhouse curiosity in 1825
(Anon. 1849), but has never flowered or fruited in Europe. MacMillan (1908)
stated that a few young trees were once grown in Syon House Gardens near
London, and were apparently presented as a gift to Queen Victoria. The 29th
edition of the Official Guide to the Royal Botanic Gardens and Arboretum, Kew
Gardens (1885) lists durian as part of the collection.
According to Soegeng-Reksodihardjo (1962), the first trees to fruit
outside Asia were grown in Dominica from seedlings shipped from Kew Gardens
in 1884. A tree introduced to the botanic gardens at St. Aroment in Dominica (15o
N) grew vigorously and fruited after about 10 years (Anon. 1894).
Durian has been grown in several locations in and about India. Firminger
and Burns (1918) gave the following diagnosis of attempts to grow durian in
Calcutta: ‘they have never risen to more than 1 metre in height, when they have
uniformly died off, the climate of that latitude being quite unsuited to them’.
However, Firminger also recorded that trees grown from seed had reached
heights of 2 metres after three years at the Burliar Experimental Gardens at the
base of Nilgiri mountains at an elevation of 760 m in southern India (11o N). Some
research on vegetative reproduction by grafting has since been carried out at
Burliar (Khan and Sambashiva Rao 1952). Durian has also been introduced into
Port Blair in the Andaman Islands (12o N) (Parkinson 1923).
Durio — A Bibliographic Review 76
There have been some attempts to introduce durian into the West
Indies and the Americas. Durian has been planted at the Botanical Gardens in
Trinidad (Pascoe 1882); however, ‘only one plant exists in the St. Clair
Experiment Station...It does not seem readily amenable to the Trinidad conditions
of climate’ (Freeman and Williams 1928). Bailey (1914) stated that durian has
been successfully introduced into Jamaica, however, I have been unable to
locate any other information or confirmation regarding this. Baker (1969)
reported a single mature tree growing in the botanical gardens at Lancetilla,
in the Honduras.
A few trees are found in Hawaii, and many bearing trees in Zanzibar
(Morton 1987). Durian is mentioned in Neal’s ‘In Gardens of Hawaii’ (1965)
and a small germplasm collection exists in Hilo (Bettencourt et al. 1992).
7
The species to which Gardner refers is probably what is now termed Cullenia ceylanica
(Gardn.) K. Schum. (see Table 1).
Durio — A Bibliographic Review 77
here and help me hunt; there must be a dead rat in my room but I can’t find
it’ — Popenoe (1956).
Clones
Clonal selection and hybridization : To date, well over 100 different Malaysian
clones of durian have been registered (Lim 1990), which are distinguished
mainly by fruit characteristics; probably more than 200 Thai durian cultivars
are recognized (Hiranpradit et al. 1987, 1992) (300 according to Malo and
Martin 1980a,b). Polchart (1952) tabulated 129 local varieties from the Dhonburi
area of Thailand alone. Fruits of the more desirable clones are sought after
and fetch a higher price in the market place (Fig. 3). Polprasid (1981) described
the use of local durian contests in Thailand to make known local cultivars and
maximize accessions of good durian germplasm. Thai durians have been divided
into six groups based on fruit morphology (Hiranpradit et al. 1987); a detailed
analysis of leaf, aril and spine morphology and fruit shape were also provided.
An itemized list of criteria for assessment of varietal desirability has been
established to aid the identification and collection of new clones (Hiranpradit
et al. 1992).
Despite the numerous Thai cultivars that have been identified, only
four (Chanee, Kradum, Mon Thong, Kan Yao) are grown on a commercial
scale (Subhadrabandhu 1993). Clones or varieties of other durian species
have not been produced or characterized. Clonal selection of durian began as
early as 1922 (Hasan and Yaacob 1986). Unfortunately, clones with very high
quality fruits (e.g. D2) are often not high yielding, while those which are
prolific yielders (e.g. D24) have lower quality fruits (Hassan, n.d.). Hassan
(n.d.) has suggested a mixture of clones is good practice in a durian orchard
to compensate for good clones being sparse bearers, and for the fact that
many durian clones are self-incompatible. The following clonal mixture is
suggested: 60% D24, 25% D16, 5% each of D10, D8, D2 (Hassan n.d.).
Kwok-Kong (1974) suggests essentially the same ratio with the substitution of
5% D7 for 5% D10. Two planting systems described by Zainal Abidin (1991b)
make use of a clonal mixture of 50% D24, 30% D99, 20% D98/D114. A row
planting system and some data on the growth and productivity of several
different durian clones grown at the experimental orchard at the Universiti
Pertanian Malaysia are given by Yaacob et al. (1978). Efficiency indices of
several individuals of three clones have been published (Hasan and Yaacob
1986).
Figure 3. Durian fruits of clone D2 and D24 for sale by the street side in
Kuala Lumpur. Durians of known and desirable clonal origin fetch
a much higher price in the marketplace.
Durio — A Bibliographic Review 79
Figure 4. The development of a durian fruit (Clone D8). The ovary undergoes
little expansion in the first month of development.
A) A fruit at 56 DAP (days after pollination), approximately 9 cm
in length. The bases of the peltate scales of the ovary have developed
into spines.
B) A fruit at 63 DAP. The fruit is approximately 10 cm in length
and shows little change from 56 DAP.
C) A fruit at 88 DAP. The fruit is now 19 cm in length and the
spines are more fully developed.
D) A fruit at 95 DAP. The fruit is now 25 cm in length and little
change occurs before the fruit abscinds at approximately 100
DAP.
Durio — A Bibliographic Review 80
established; in Malaysia between 1986 and 1989 over 800 000 clonally propagated
durian trees were produced for planting (Ali 1993). This increase has led to
a need to positively identify clonal material as mistakes have expensive and
very long-term consequences. Watson (1993), for example, reported that problems
in the introduction of durian as a crop in Australia were due to the importation
of misidentified clones.
Durian clones and varieties have been selected mainly for desirable
fruit characteristics. Thus, durian varieties can often be identified by differences
in fruit morphology. As mentioned previously, the shape of the spines has
been described for several Thai varieties (Hiranpradit et al. 1992). Hiranpradit
et al. (1987) claim that fruit shape and spine morphology are useful for
characterizing durians into groups, and both are highly heritable characteristics.
However, given the vast number of clones/varieties now recognized, it seems
rather unlikely that all fruits could be sufficiently identified from the shape of
their spines.
been published (Salma 1993). This preliminary work showed that 5 durian
clones used in the study could be distinguished by this technique; the isozyme
patterns studied were those of acid phosphatase, alkaline phosphatase and
peroxidase.
Seeds : Direct sowing of seeds in the field is not recommended due to damage
from rodents (Anon. 1953a). Shaharuddin (1979) has also reported large losses
of germinated seeds due to rodents which eat the epicotyls. Although seedlings
usually cannot survive such injuries, De Vogel recorded an instance of a
seedling of D. zibethinus in which a ring of buds developed from a region
between the cortex and the stele on a seedling whose epicotyl (including the
top of the hypocotyl) had been bitten off (De Vogel 1980). If planted in a
seedbed, Coronel et al. (1983) recommended that seeds be planted 1 cm deep
with a spacing of 4-6 cm between them. Coronel et al. (1983) also recommended
the coating of the seeds with fungicide.
Root pruning : Very little information on the roots of durian was available
until very recently. ‘The tree is very impatient of disturbance of its roots’,
(Anon. 1935) however, another early report stated that cutting the roots of
Durio — A Bibliographic Review 82
durian seedlings with a spade promotes the production of a fibrous root system
reducing loss during transplanting (Feilden and Garner 1936). More recently,
Chong (1985) suggested that the roots of durian should not be pruned to
reduce circling or kinking (as commonly occurs when young trees are grown
in polybags), as the trees are very sensitive to such treatment. This conflict
has very recently been resolved by Ghani (1992a,b). This empirical study
demonstrated that root pruning increased the growth of a fibrous root system,
contributed to increased overall growth and stem enlargement, and increased
survival rate of transplanted trees.
What all nutrient removal studies have shown is that K is much more
important than N for durians, being the major element removed by the fruits
and by the tree itself, the amount of P being small in comparison. This is
supported by the study of Jamil (1992c) who examined the effects of N, P and
K on young durian trees. Increasing N was found to have no visible effect on
plant form, increased P increased the tree height, while increased K greatly
affected tree form. Thus, of the former cited fertilizer suggestions, those
recommending high potassium formulations are the most valid.
inclines up to 35o (Hassan n.d.). Maas et al. (1979) stated that durian is
suitable for growth on slopes up to 25o. A study of moisture requirements
of young trees on steep (30 o) and low (19o) slopes demonstrated that trees
on steeper slopes fared less well, even with identical rainfall, to those on
less steep slopes, due to the lower soil moisture reserve (Yaacob 1992).
Poorer growth on steep slopes can be alleviated by mulch and irrigation
(Yaacob 1992).
with sides of 75 cm, but larger holes are more advantageous (Hassan n.d.).
The removed soil should be mixed with 12 kg of manure and replaced. The
young trees can be planted one week after the hole has been prepared (Hassan
n.d.). It has been suggested that organic manures may facilitate infection by
P. palmivora (Watson 1984).
A study on the most suitable size material for planting in the field
revealed that budded material of clone D24 with a stem circumference of
3 cm had 66-77% survival after one year. Saplings with smaller stem
circumferences had survival rates as low as 29% (Ghani 1988). Some clones
(D24 in particular) are apparently difficult to establish in orchards (Ghani
1988).
Intercropping : Young durian trees are sensitive to strong sunlight and should,
therefore, be intercropped with other plants to provide shade (Hassan n.d.).
The major concern for intercropping is that the intercrop does not harbour
Phytophthora (Coronel 1986). It is, of course, an additional benefit if the
intercrop is itself economically valuable. Bananas have been recommended
for this purpose (Hassan n.d.; Hashim et al. 1985; Mohamad Idris 1987).
Coronel et al. (1983) stated that newly planted trees are shaded with bananas
for the first three or four years. Pineapples are also used for intercropping
(Coronel et al. 1983). Gliricidia and guava have also been recommended for
intercropping (Mohamad Idris 1987). Osman and Basri (1987) recommended
intercropping with cocoa. Conversely, durian trees are used as shade trees for
young cocoa trees (Anon. 1991a). A planting scheme for intercropping durian,
cocoa and Gliricidia is presented by Jelani et al. (1992). It has been suggested
that intercropping durian with cocoa can lead to increased incidence of serious
pathogens such as Phytophthora and Rhizoctonia. Nawi and Mohd. (1991)
have studied this claim and conclude that it is largely unfounded, any small
increase in outbreaks of disease that may be associated with this intercropping
can be readily controlled by improving crop management practices. Coronel
Durio — A Bibliographic Review 87
et al. (1983) recommended that papaya and coconut should not be used to
intercrop durian for reasons of disease control.
Post-harvest Technology
Information regarding the harvesting and post-harvest processing of
durian fruits is of great importance as the two factors that seriously limit the
durian fruit development as a crop are its smell and its short shelf-life; both
of these factors can be controlled or affected by harvest and post-harvest
practices. Most of the post-harvest technology research has originated in Thailand.
Although Thailand has made great strides in this area, much of the published
research is difficult to obtain and is written in the Thai language, which
undoubtedly limits its exploitation elsewhere. Thai research has produced
valuable information in numerous areas, such as harvesting, local transport
and preparation of fruits for market, methods to overcome the difficulties of
export peculiar to this fruit, and detailed examination of the physiology of
ripening. All of this research has provided insight into how durian's two most
limiting properties can be overcome. Experiments in the physiology of fruit
ripening, in particular, suggest methods to manipulate ripening which offer
the prospects of greatly extending the shelf-life and limiting the undesirable
nature of its smell.
Durian fruits generally fall from the trees at night (Teo 1991). In
Malaysia, fruits are normally collected after they fall (Watson 1984; Mohamed
1990) as it is believed that harvesting the fruits before they are mature affects
the flavour (Nanthachai et al. 1994). In Thailand, fruits are detached from the
tree just before maturity (Watson 1984) and then allowed to ripen. A recent
study by Pauziah et al. (1992) demonstrated that Malaysian D24 durians
harvested at 105 and 110 days after anthesis and allowed to ripen had the
same organoleptic properties as mature fallen fruits; however, fruits harvested
too early (100 DAP) did not ripen properly. Hand harvesting immature D24
fruits increased the shelf-life to 9-11 days from that of only 3-4 days for
fallen fruits (Pauziah et al. 1990,1992). Similar conclusions were reached in
a study in the Philippines (Pascua and Cantila 1991). Thus, knowledge of the
Durio — A Bibliographic Review 88
ripening process, and the exact time at which to harvest immature fruits of
each particular clone or variety can be of great value in extending the shelf-
life of the fruit, and yet not affect fruit quality.
at -23oC. After longer storage, arils started losing flavour (Anon. 1960). It is
reported that preliminary experiments in freezing durian arils in polyethylene
bags at -22oC succeeded fairly well. Praditdoung (1986) reported that the
shelf-life of intact durian arils (containing seeds) could be extended up to 30
days when wrapped in low density polyethylene and stored at 4oC. Moleeratanond
et al. (1990) demonstrated that the quality of durian arils wrapped in plastic
and kept at 2oC could be maintained for 48 days for Chanee durians, and 30
days for Mon Thong. Numerous physical data (weight loss, CO2 buildup, etc.)
were tabulated for different treatments. Romphophak and Palakul (1990), who
studied cold storage of whole fruits, provided results somewhat contrary to
those just described. Using Chanee durians kept at 5oC, either untreated or
treated with calcium carbide, they found the pericarp of cold stored durians
showed signs of chilling injury after just one week. Soluble solids in cold
stored fruits remained low and did not rise even 3 days after removal to room
temperature. Durian pulp remained firm while in cold storage, and became
soft upon removal to room temperature. Thus, aril softening and increase in
soluble solids are apparently governed by separate processes. More importantly,
they reported that the palatability, scored by a panel of 6 members, was much
lower for fruits stored for even 1 week than for unrefrigerated controls. No
increase in palatability of refrigerated fruits occurred even after returning the
fruits to room temperature for up to 3 days.
packaging method for intact ripe durian fruits. Corrugated fibreboard boxes
(5.24 mm thick) with an area of ventilation of 2.5% were selected as the
best type of export shipping container (Paklamjeak et al. 1986). Furthermore,
Paklamjeak et al. (1986) showed that the belief that inclusion of a few basil
leaves in the packing container will eliminate or absorb the odour was false,
however, no other consideration was given in their study to smell containment
properties of a desirable container. The design and structural features of
durian export packaging containers is further elaborated by Paklamjeak et
al. (1988, 1989).
Durian fruits typically split open as they ripen. This splitting is due
to the development of an abscission zone. Very little is known about structure
Durio — A Bibliographic Review 91
of the abscission zone, however, the abscission zone cells are known to have
tannin deposits (Sriyook et al. 1994). Low humidity is known to stimulate
dehiscence (Ketsa and Pangkool 1994). Durians loose a lot of moisture after
harvest. In one study, moisture content was shown to decrease by 21% after
9 days of storage at 30oC and 70% relative humidity, although moisture loss
was lower when fruits were stored at higher humidities (Ketsa and Pangkool
1994). The majority of the moisture lost originates from the pericarp, no
significant drop in moisture content of the arils was found even after 5 days
at 75% RH (Ketsa and Pangkool 1994).
Plant growth regulators : CO2 and ethylene show a parallel increase during
ripening of durian fruits as shown by Tongdee et al. (1990) in their study on
the effect of O2 and CO2 on ripening. Furthermore, CO2 and ethylene production
remain high after the climacteric (Tongdee et al. 1989a). A high ethylene
level at the time of harvest (1.4 ppm) in Chanee durians indicates that ethylene
actually accumulates before fruit abscission, at least in fruits of this variety
(Tongdee et al. 1989a).
ripening of the arils stimulates the ripening of the husk in intact fruits (Booncherm
and Siriphanich 1991).
Forestry Aspects
Coelostegia griffithii Mast. : The weight of the wood has been recorded as
713 kg/m3 by Ridley (1901, 1903), and 705 kg/m3 at 15% moisture by Burgess
(1966). The bark of this tree is used to tan nets. The wood is hard, flexible
and durable (Ridley 1901, 1903).
Durio carinatus Mast. : The bark of this species is apparently used for roofing
(Uphof 1968; Usher 1974). An examination of charcoal, charcoal briquets and
alcohol production from different types of wood waste of D. carinatus is
presented by Syachri (1983).
The descriptive measurements from Chu (1969) are: air dry density-
average 630 kg/m2 range 551-700 kg/m3; fibre length - average 1.65 mm,
range 1.55-1.79mm; fibre average maximal tangential diameter ave. 32.63
µm, range 30.45-34.10 µm; fibre average maximal tangential lumen ave. 25.71
µm, range 24.26-28.32 µm; fibre average maximal tangential wall thickness
ave. 7.25 µm, range 5.89-9.08 µm.
Durio dulcis Becc. : An analysis of wood from this species is given by Sudradjat
(1980). The wood is listed as having a specific gravity of 0.73. Cellulose
accounts for 50.9% of the oven dry weight of the wood, 36.7% is accounted
for by lignin, 14.6% pentosan and 1% ash. Cockrell (1942) listed the specific
gravity of the wood of D. conicus Becc. [=D. dulcis Becc. sensu Kostermans
and Reksodihardjo 1958] as 0.60.
Durio ?lowianus Scort. ex King : The weight of wood from this species was
listed as 657 kg/m3 air dry (Desch 1941).
Durio malaccensis Planch. ex Mast. : The weight of wood from this species
was listed as 705 kg/m3 air dry (Desch 1941).
Durio ?oblongus Mast. : The weight of wood from this species was listed as
657 kg/m3 air dry (Desch 1941).
The weight of the wood was listed at 753 kg/m3 air dry by Desch
(1941) and 755 kg/m3 by Ridley (1901, 1903). A value of 610 kg/m3 at 15%
moisture was listed as the mean weight of 10 specimens by Burgess (1966).
Durio — A Bibliographic Review 96
Durio zibethinus L. : The wood of this species is a dull red brown colour with
a hard grain, yielding an uneven rough surface. It is liable to warp and unsuitable
for export (Howard 1948). The wood is straight grained and moderately heavy
(Keith 1947). The wood has a specific gravity of 0.42 (Cockrell 1942). The
weight is listed as 481 kg/m3 (air-dried) by Keith (1947); 545 kg/m3 air dry
by Desch (1941) and Howard (1948); 570 kg/m3 at 15% moisture by Burgess
(1966); and 645 kg/m3 by Ridley (1901, 1903). The wood is suitable for
general construction (Keith 1947).
Uses of ‘Durian’ timber : Durian wood is used for making clogs in Sarawak
(Foxworthy 1921; Thomas 1952, 1979; Anon. 1964). It is also used for light
construction, cheaper grades of furniture (Thomas 1952, 1979; Anon. 1964)
and temporary construction work (Foxworthy 1909). Durian wood is
recommended for non-impact tool handles (Lim 1988) and cigar boxes
(Martawijaya and Kartasujana 1981). Durian wood makes satisfactory plywood
(Thomas 1952, 1979). Studies of the gluability of durian plywood have been
conducted by Tsai (1975); the wettability of wood veneers of durian with
different adhesives has also been studied (Wang 1975), as has the bonding
Durio — A Bibliographic Review 97
Figure 5. The inside of mature fruit of D. zibethinus. Two of the five locules
are visible. The seeds are covered within the fleshy yellow arils
which surround them and fill the locules.
Durio — A Bibliographic Review 98
strengths of different adhesives (Wang 1977; Taki 1986). Tensile tests have
been conducted on plywood made from durian (Kitamura et al. 1982).
Wood anatomy : The wood anatomy of Durio has been examined in some
detail by several authors (Bargagli-Petrucci 1904; Moll and Janssonius 1906;
Cockrell 1942; Menon 1959). The wood anatomy of D. lowianus is described
in a Japanese paper (Anon. 1966). According to Menon (1959), there is silica
in the wood parenchyma of B. griffithii Mast. [=D. griffithii (Mast.) Bakh.
sensu Kostermans 1958b] and C. griffithii Benth.; D. grandiflorus is also
siliceous (Burgess 1966). No silica is present in D. oxleyanus, D. testudinarum,
D. zibethinus or Neesia altissima. There are crystals in chambered parenchyma
strands in all ‘durian’ save B. griffithii Mast. [=D. griffithii (Mast.) Bakh.
sensu Kostermans 1958b] (Menon 1959). Vertical traumatic intercellular canals
are present (Metcalfe and Chalk 1957). Mucilage cavities are present in the
pith of Durio, Coelostegia and Boschia (Solereder 1908).
Growth rings are distinct, and are delineated by denser fibrous tissue
at the outer margin (Cockrell 1942). The sapwood of durian is white or pale
yellow-brown, while the heartwood is brown to deep brown (Anon. 1964).
Both uni- and multi-seriate rays are present in the wood. Measurements
of medullary ray elements from several durian species are tabulated by Bargagli-
Petrucci (1904). The xylem rays of Durio are characterised by the presence
of tile cells (rows of narrow upright cells that resemble tile work when viewed
in radial section). Tile cells typically occur in certain members of the Malvales.
These tile cells are grouped into two types. The first type are the Durio type
tile cells, these are about the same height as the procumbent ray initials, as
Durio — A Bibliographic Review 100
is typical in the genus Durio and 17 other genera in the Malvales (Manchester
and Miller 1978). Additionally, there exists the Pterospermum type of tile
cell, in which the tile cells are two to several times higher than the procumbent
initial cells. This type of tile cell is found in approximately 30 Malvalceous
genera (Manchester and Miller 1978). The development of Durio type tile
cells has been described by Chattaway (1933). Resin is abundant in procumbent
cells of durian, but rare in tile cells. The tile cells often contain crystals
(Chattaway 1933). Durio type tile cells are found in all members of the
Durioneae except Camptostemon and Maxwellia (Metcalfe and Chalk 1957).
A study of fossilized eocene wood revealed the earliest known existence of
tile cells, these being of the Pterospermum, or an intermediate Pterospermum-
Durio type. No known fossil woods with Durio type tile cells are known,
however, based on the mode of development, Manchester and Miller (1978)
speculated that the Pterospermum type may have arisen from the Durio type.
Bacteria
Enterobacter sp. -stem canker Liu (1977b); Singh (1980) Flavobacterium sp.
-stem canker Liu (1977b); Singh (1980); Pseudomonas sp. -stem canker Liu
(1977b); Singh (1980).
Durio — A Bibliographic Review 101
Fungi
Cercospora sp. -leaf spot, Chandrasrikul (1962), Giatgong (1980), Singh (1980),
Williams and Liu (1976),
Corticium solani (Prill & Delacr.) Bourd. & Galz. -leaf spot and dieback,
Singh (1973, 1980); Turner (1963; 1971).
Corynespora cassiicola (Berk. & Curt.) Wei. - associated with leaf blotch,
Anon. (1967); Singh (1980); Williams and Liu (1976).
Homostegia durionis Rac. -black and brown leaf spots, Coronel et al. (1983);
Soegeng-Reksodihardjo (1962); Subhadrabandhu et al. (1991).
Leptoxyphium sp. -black crusty growths on leaves and twigs, Lim (1989).
Marasmiellus scandens (Mass.) Dennis & Reid -thread blight, Singh (1980),
Turner (1971).
Meliola durionis Hansf. -black mildew, sooty mould, Hansford (1956); Johnston
(1960); Lim (1989); Nik Masdek et al. (1991); Singh (1973, 1980); Turner
(1961, 1971).
Nectria sp. -associated with bark rot, Anon. (1954); Nectria haematococca
Berk. & Br. -stem rot, Johnston (1960); Singh (1973, 1980); Nectria ochroleuca
(Schw.) Berk. -stem rot, Johnston (1960); Singh (1973, 1980).
Phomopsis sp. -leaf spot, Chandrasrikul (1962); Giatgong (1980); Singh (1973,
1980); Soegeng-Reksodihardjo (1962); Thompson and Lim (1965); Phomopsis
durionis Syd. -leaf spot of seedlings, branch and stem necrosis, Liu (1977b);
Nik Masdek et al. (1991); Singh (1973; 1980); Weber (1973).
Phragmocapnias betle (Sydow & Butler) Theissen & Sydow emend. Reynolds.
Durio — A Bibliographic Review 104
Phyllosticta sp. -leaf spot, seedling rim blight, Anon. (1967); Liu (1977b);
Singh (1973; 1980); Thompson (1939); Tidbury (1976); Williams and Liu
(1976); Phyllosticta durionis A. Zimmermann -leaf spot, rim blight, Chandrasrikul
(1962); Coronel et al. (1983); Giatgong (1980); Hassan (n.d.); Soegeng-
Reksodihardjo (1962); Subhadrabandhu et al. (1991); Weber (1973).
Polychaeton sp. -leaf mould and black crust of fruit, Lim (1989).
Pyrenochaeta sp. -leaf spot, Johnston (1960); Singh (1973, 1980); Williams
and Liu (1976).
Durio — A Bibliographic Review 105
Pythium sp. -root rot, Coronel (1986); Lee et al. (1993); Ng and Choong
(1991); Nik Masdek et al. (1991); Pythium complectans Braun [synonym of
P. vexans de Bary] -root disease, patch canker., Belgrave (1939); Coronel et
al. (1983); Johnston (1959); Meon and Varghese (1986); Pawakul and
Chittanawasarn (1977a; 1977b); Roger (1951); Singh (1973, 1980); Soegeng-
Reksodihardjo (1962); Thompson (1938; 1939); Watson (1984); Weber (1973);
Pythium vexans de Bary [see also P complectans which is a synonym], Ling
(1991); Shaji et al. (1993)
Rhizoctonia sp. -leaf fall, leaf blight, Chandrasrikul (1962); Giatgong (1980);
Vichirananda (1983); Rhizoctonia solani Kuhn. -leaf blight, Anon. (1986b);
Lim et al. (1987); Ng et al. (1986); Ng and Choong (1991); Nik Masdek et
al. (1991); Shaji et al. (1993); Singh (1973; 1980).
Rigidoporus lignosus (Klotzsch) Imazeki [see also the synonym Fomes lignosus]
-white root disease, Ng and Choong (1991); Nik Masdek et al. (1991); Shaji
et al. (1993); Singh (1980); Williams and Liu (1976)
Trichomerium grandisporum (Ellis & Martin) Bat. & Cif. -sooty mould of
leaves, petioles and twigs, Lim (1989).
Trichopeltheca asiatica Bat., Costa & Cif. -black film on leaves, Lim (1989).
Durio — A Bibliographic Review 106
Lichens
Algae
Cephaleuros virescens Kunze. -red rust, Nik Masdek et al. (1991); Singh
(1980); Vichirananda (1983); Williams and Liu (1976); Phycopeltis sp. -largely
harmless epiphyte, Lim (1990); Pleurococcus nagelii -largely harmless epiphyte,
Lim (1990); Trebouxia sp. -largely harmless epiphyte, Lim (1990); Trentepohlia
sp., Long et al. (1991); Trentepohlia aurea (L.) Martius, Lim and Sijam
(1989b); Trentepohlia monile De Wildeman [=Physolinum monilis (De Wildeman)
Prinz. =T. moniliformis Karsten], Lim and Sijam (1989b); Trentepohlia arborum
(C.Ag.) Hariot [=T. bisporangiata Karsten.], Lim and Sijam (1989b).
Ferns
Angiosperms
Insects
Asterolecanium sp. -leaf scales, Anon. (1959a); Chuan et al. (1981); McIntosh
(1951); Yunus and Hua (1980); Asterolecanium ungulata Russel, Lim et al.
(1991); Long et al. (1991); Yunus and Hua (1980).
Batocera gultata Vollenh. -stem borer, Nik Masdek et al. (1991); Shaji et al.
(1993); Yunus and Hua (1980).
Carpophilus sp. -sap beetle, Lee et al. (1994); Carpophilus floveicollis -sap
beetle, Lee et al. (1994).
Daphnusa ocellaris Walk. -Hawk moth which eats durian leaves, Ramasamy
(1980).
Hypomeces squamosus F., Shaji et al. (1993); Yunus and Hua (1980).
Microtermes pallidus (Haviland), Nik Masdek et al. (1991); Yunus and Hua
(1980).
Monolepta bifasciata Hornst., Shaji et al. (1993); Yunus and Hua (1980).
Orgyia postica Walk., Yunus and Hua (1980); Orgyia turbata Butl., Yunus
and Hua (1980).
Phostria nicoalis Wlk., Yunus and Hua (1980); Phostria xipharesalis Wlk.,
Durio — A Bibliographic Review 109
Platypus cupulatus Chap., Nik Masdek et al. (1991); Yunus and Hua (1980).
Sylepta derogata F., Yunus and Hua (1980); Sylepta bipunctalis Warr., Yunus
and Hua (1980).
Synanthedon sp., Nik Masdek et al. (1991); Yunus and Hua (1980).
Tetranychus fijiensis Hirst., Lee et al. (1994); Yunus and Hua (1980).
Tirathaba mundella Walk., Yunus and Hua (1980); Tirathaba ruptilinea Hamps.,
Frannsen (1936).
Tonica terrasella Wlk. -fruit borer, Lee et al. (1994); Mohd. Shamsudin (1991);
Mohd. Shamsudin and Norsiah (1990); Mohd. Shamsudin (1992); Nik Masdek
et al. (1991); Zainal Abidin et al. (1992).
Sch., Yunus and Hua (1980); Xyleborus ferrugineus (F.), Yunus and Hua
(1980); Xyleborus formicatus Eichh., Boonyong (1983); Xyleborus similis Ferr.,
Yunus and Hua (1980); Xyleborus testsceus Wlk [=X. perforans Wall.], Yunus
and Hua (1980).
Zeuzera sp. -borers., Polchart (1952); Zeuzera coffeae Nietn. -stem borer,
Coronel et al. (1983), Nik Masdek et al. (1991), Yunus and Hua (1980).
Nematodes
Helicotylenchus sp., Fong and Poh (1970); Sidam and Yuen (1992); Singh
(1973).
Macroposthonia sp., Sidam and Yuen (1992).
Meloidogyne sp., Sidam and Yuen (1992).
Pratylenchus sp., Sidam and Yuen (1992); Pratylenchus coffeae
(Zimmerman)Filipjev & Schuurmans Stekhoven -hypocotyl rot, Giatgong (1980).
Radopholus sp., Fong and Poh (1970); Singh (1973).
Rotylenchulus reniformis Lin. & Ol., Sidam and Yuen (1992).
Tylenchorhynchus sp., Sidam and Yuen (1992).
Xiphinema sp. -attack roots, Ng and Choong (1991); Sidam and Yuen (1992).
Other animals
Although durian seeds are viable and trees have historically been grown
from seed, durians are now preferably propagated asexually by grafting, seeds
being used for the production of rootstocks. Seeds of the cultivar Chanee are
used to produce rootstocks for grafting in Thailand (Subhadrabandhu et al.
1991). The major problems with direct sowing of seeds are that the genetic
makeup of the trees is unknown, and the emerging seedlings are susceptible
to predation by rodents. Seeds are now generally grown in a nursery in polybags
to produce material for grafting (Hassan n.d.).
Consequently, more grafted trees can be planted per hectare of land and the
fruits are produced closer to the ground. Thirdly, grafted trees start bearing
fruits at an earlier age than do trees raised from seed.
The disadvantages of vegetatively propagated material are likewise
three-fold. Vegetative propagation is much more labour intensive, different
grafting methods have varying degrees of success, and durian trees are sensitive
to transplanting; budded durian trees frequently suffer 50% mortality (Feilden
and Garner 1936). Thus, it is preferable to keep trees in the nursery for
extended periods after grafting to increase the success of transplanting. Because
of the great advantages to be gained by asexual propagation, much research
has been conducted in this aspect of durian cultivation.
Double root system : ‘Three legged’ durian trees are created in Thailand
(Boonbongkarn 1960). Using this technique, the stems of a few seedlings are
grafted onto a more mature tree. This is done after the tree has reached
1-1.5 m in height. The resultant tree has more than one root system, is better
able to obtain food and water, and can withstand stronger winds (Boonbongkarn
1960). Old durian trees with declining productivity can apparently be revitalised
by grafting a young shoot onto the trunk, and then removing the original stem
(Boonbongkarn 1956). Ahmad and Ghani (1990) have noted that this technique
is occasionally used on D. zibethinus to induce rapid flowering.
Approach grafting : Success with approach grafting of durian trees has been
reported using 6-8 months old seedlings (Chua and Teoh 1973). Up to 100%
success was reported by Chua and Yong (1978), however, success of graft
Durio — A Bibliographic Review 113
was higher using older (8-10 months) rootstocks, and an extended time before
cutback. Baga Kalie (1979) described a technique for simple splice approach
grafting of durian. Baga Kalie and Anwarudin (1980) compared results of
simple splice approach grafting and tongued approach grafting of durian;
these two techniques were found not to differ significantly in success rate.
The highest rate of success was achieved using 8.5 months old rootstocks. An
approach grafting technique for grafting seedlings onto branches of mature
trees is presented by Zainal Abidin et al. (1991). In Thailand, bagged rootstocks
are also grafted onto branches of mature trees by inarching (suckle grafting)
(Subhadrabandhu et al. 1991).
Top grafting : The effects of scion length for top grafting has been investigated
using four Thai cultivars of durian (Anwarudin and Sunarjono 1987). Scions
of 2, 3 and 4 nodes length were tested; no significant difference in graft take
was found. Some durian varieties were found to top graft more easily than
others. Rootstocks of different ages have also been tested to discover the most
suitable age. No significant difference in suitability was found in rootstocks
aged between 1.5 and 3 months (Anwarudin et al. 1987).
Naik (1949) stated that inarching experiments met with 50% success,
but there was no success in shield budding; however, he did not describe his
shield budding technique. Wester (1918) provided the following instructions
regarding shield budding of durian: ‘Use non-petioled, fairly mature, but not
old and hard budwood; cut the bud 2.5 to 3.5 cm long; age of stock at point
of insertion of bud unimportant’.
branches. It is important to note that buds will fail to unite if they become wet
(Sankhariksha 1968; Coronel et al. 1983).
Bud grafting experiments with durian have showed that a high percentage
success can be obtained. Younger rootstocks are preferable (in this case 4
months) (Chua and Yong 1978). Coronel (1986) stated that patch budding is
the least popular method used in the Philippines due to the complexity of the
technique, and the need for more mature plant material. Commercial bud
grafting of durian has been practised in Malaysia since at least 1954 (Dickinson
1959); according to Hassan (n.d.), bud grafting is the most common form of
durian propagation practised in Malaysia.
Very little work on the tissue culture of durian has been published.
Some success with shoot tip culture of D. lowianus has been reported. Although
durian tissue can be made to callus, no report of its regeneration has been
published.
Grafting to other species : Experiments have been carried out at the Burliar
Fruit Station to graft D. zibethinus scions onto Cullenia excelsa rootstocks
(Anon. 1953b). Grafting onto Cullenia excelsa rootstocks in India has been
reported by Sundararaj et al. (1970). Tidbury (1976) tabulated results of grafting
experiments onto rootstocks of this species at the Burliar Research Station.
These results showed that approach grafts to C. excelsa rootstocks averaged
much greater growth than did grafts to D. zibethinus rootstocks. Grafting to
C. excelsa also contributed to earlier flowering. Unfortunately, no record of
actual success rates of grafts to C. excelsa rootstocks were uncovered in the
available literature.
Durio — A Bibliographic Review 116
especially since such grafted trees would need to be planted in large numbers
to significantly overcome the economic effects of the disease. The broken
resistance of D. malaccensis stands as an example of the futility of this simplistic
approach (Pupipat 1984). Grafting to other species could perhaps become a
useful tool to produce trees of smaller stature, trees which bear fruit at an
earlier age, and possibly grafted seedlings with a greater survival rate upon
transplanting. Future research on inter-specific grafts would do well to concentrate
on these objectives.
Significant genetic erosion within the genus Durio has already been
reported (Sastrapradja 1975), especially in Malaysia, Thailand and Indonesia,
where clonal varieties have become popular. Hasan (1989) stated that within
Malaysia, durians should be the first priority in development. Karim (1992)
suggested that different flowering seasons of different clones should be
exploited on a national plantation scheme to spread the durian season. This
idea is thoroughly discussed by Zainal Abidin et al. (1986) (see section
on manipulating seasonality). Karim (1992) also suggested that research
is needed to reduce the gestation time of the fruit. Pietrzyk (1992) has
stated the need for the creation of cDNA libraries for durian and other
Malaysian fruits.
Durio — A Bibliographic Review 120
as it is a crop that has undergone very little domestication and one which has
a large potential for improvement. The genetic resources of the common
durian are threatened, to some extent, apart from deforestation, due to the new
'improved' clones themselves. The proliferation of popular varieties has led to
considerable genetic erosion as less popular non-clonal trees are replaced
(Sastrapradja 1975). This is a particular problem in Indonesia, Malaysia, Thailand
and Vietnam (Smith et al. 1992).
The wild species of Durio present another and perhaps larger challenge
for genetic conservation. D. zibethinus faces the dilution of its gene pool by
the large scale planting of genetically identical clones. By comparison, all
of the wild species are quite rare, some are in danger of extinction and a few
may already be extinct. It is claimed that a D. macrantha tree growing in
Bogor (a newly described species with edible fruit) may be one of the last
living specimens (Anon 1992). Smith et al. (1992) stated that several wild
species are threatened by logging. Although admittedly not much is known
about the genetic diversity of most of the wild species, erosion has been
documented for D. kutejensis (Sastrapradja 1973b). The genetic resources of
the wild species are equally, if not more, valuable than those of D. zibethinus
for three reasons. Firstly, some of these species do have potential as fruit
tree crops themselves. Although not grown anywhere near the scale of D.
zibethinus, several other species are cultivated. D. kutejensis is cultivated in
the eastern part of Kalimantan and D. oxleyanus is in rare instances cultivated
in some Bornean villages (Soegeng-Reksodihardjo 1962). At least a dozen
of the known species have edible fruits (see section on edibility), although
the majority of these have never been cultivated. Some of the these wild
fruits may actually be superior in quality to those of many of the clones of
D. zibethinus. Secondly, there is evidence that some of the wild species may
have higher resistance to some of the major durian pathogens (Phytophthora
in particular). Hamilton (1975) has already called for the investigation of
resistant rootstocks to P. palmivora from other durian species. There is
interest in not only using wild Durio species as rootstocks for D. zibethinus
(as Phytophthora is a root disease), but in producing true hybrid trees.
Thirdly, the possibility of producing hybrid trees opens the door to improvement
possibilities far beyond resistance to pathogens. Several Durio species are
so closely related to D. zibethinus, they most likely do, or have already been
shown to hybridize with it. Possibilities thus exist for altering the size of the
tree, time of flowering, reducing the age to first flowering, affecting seasonality,
increasing yield, improving the flavour, smell, colour, texture and size of
the fruits.
Bogor (National ? ? 2
Biological Institute)
(Contd....)
Durio — A Bibliographic Review 124
“Poets and moralists, judging from our English trees and fruits, have
thought that there existed an inverse proportion between the size of the one
and the other, so that their fall should be harmless to man. Two of the most
formidable fruits known, however, the Brazil Nut (Bertholletia) and the Durian,
Durio — A Bibliographic Review 126
grow on lofty trees, from which they both fall as soon as they are ripe, and
often wound or kill those who seek to obtain them. From this, we may learn
two things: first, not to draw conclusions from a very partial view of Nature;
and secondly, that the trees and fruits and all the varied productions of the
animal kingdoms, have not been created solely for the use and convenience
of man” — Wallace (1856)
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